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Acta Musei Moraviae, Scientiae biologicae Special issue, 98(2)

Studies in in honour of Pavel Lauterer and Jaroslav L. Stehlík

PETR KMENT, IGOR MALENOVSKÝ & JIØÍ KOLIBÁÈ (Eds.)

ISSN 1211-8788 Moravian Museum, Brno 2013 RNDr. Pavel Lauterer (*1933) was RNDr. Jaroslav L. Stehlík, CSc. (*1923) born in Brno, to a family closely inter- was born in Jihlava. Ever since his ested in natural history. He soon deve- grammar school studies in Brno and loped a passion for nature, and parti- Tøebíè, he has been interested in ento- cularly for . He studied biology mology, particularly the true bugs at the Faculty of Science at Masaryk (). He graduated from the University, Brno, going on to work bri- Faculty of Science at Masaryk Univers- efly as an entomologist and parasitolo- ity, Brno in 1950 and defended his gist at the Hygienico-epidemiological CSc. (Ph.D.) thesis at the Institute of Station in Olomouc. From 1962 until Entomology of the Czechoslovak his retirement in 2002, he was Scienti- Academy of Sciences in Prague in fic Associate and Curator at the 1968. Since 1945 he has been profes- Department of Entomology in the sionally associated with the Moravian Moravian Museum, Brno, and still Museum, Brno and was Head of the continues his work there as a retired Department of Entomology there from research associate. Most of his profes- 1948 until his retirement in 1990. sional career has been devoted to the During this time, the collections study of psyllids, , plant- flourished and the journal Acta Musei hoppers and their natural enemies. He Moraviae became established as a sci- has built up a huge collection of these entific periodical. To date, he has pub- groups and has published over 200 sci- lished over 120 scientific papers, many entific papers on their systematics and of them addressing the systematics faunistics, as well as applied entomo- and of world logy. He has also always been an ent- and and the faunistics of husiastic populariser of entomology, Czech and Slovak Heteroptera. especially among younger people. EDITORIAL

This specially-published edition of Acta Musei Moraviae, Scientiae biologicae is dedicated to two outstanding entomologists, RNDr. Jaroslav Stehlík, CSc. and RNDr. Pavel Lauterer who, in 2013, celebrated their ninetieth and eightieth birthdays respectively. Both have devoted their entire professional careers to the Hemiptera and to the Moravian Museum. In the museum, they have, for several decades, set consistently high standards of research and built up huge scientific collections in the Department of Entomology. They kept both the museum and Czech entomology in continuous contact with the international scientific community, even in the times when politics rendered this exceptional and far from easy, and when the world was far less technically interconnected than that which we take for granted today. We include 27 contributions on the taxonomy, systematics, biology and faunistics of the Heteroptera, and – insect groups close to the hearts of Jaroslav Stehlík and Pavel Lauterer – from a total of 53 authors from all over the world. In addition to the authors, many other colleagues have participated in the volume with peer-reviews of individual papers or personal memories: Charles Bartlett (University of Delaware, Newark, USA), Jérôme Constant (Royal Belgian Institute of Natural Sciences, Brussels, Belgium), Jakob Damgaard (Zoological Museum/Zoological Institute, Copenhagen, Denmark), Dmitry Dmitriev (Illinois Natural History Survey, Champaign, USA), Jowita Drohojowska (University of Silesia, Katowice, ), Sakis Drosopoulos (Athens, Greece), Murray Fletcher (Orange Agriculture Institute, Orange, Australia), Dimitri Forero (Pontificia Universidad Javeriana, Bogotá, Colombia), Paul Freytag (University of Kentucky, Lexington, USA), Ian Hodkinson (Liverpool John Moores University, Liverpool, UK), Gernot Kunz (Graz, ), Mallik Malipatil (Bioprotection, Ferntree Gully Delivery Centre, Australia), Lorèn Marchal (Muséum national d’Histoire naturelle, Paris, ), Felipe F. F. Moreira (Instituto Oswaldo Cruz, Brazil), Wolfgang Rabitsch (Umweltbundesamt, and Nature Conservation, Vienna, Austria), David Rider (North Dakota State University, Fargo, USA), Dariusz Œwierczewski (Jan D³ugosz University, Czêstochowa, Poland), and James Zahniser (Illinois Natural History Survey, Champaign, USA). Tony Long (Svinošice) sub-edited and helped to work up most of the English texts with his usual enthusiasm and close attention to detail. We thank all of these people for their contributions and, on their behalf, extend to Jaroslav Stehlík and Pavel Lauterer the “Many Happy Returns” that their years and the anniversaries merit.

Petr Kment, Igor Malenovský and Jiøí Kolibáè

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 5–10, 2013

On the occasion of the ninetieth birthday of RNDr. Jaroslav L. Stehlík, CSc.

On April 11, our dear colleague and long the mentor to Czech and Slovak heteropterology, RNDr. Jaroslav L. Stehlík, CSc., celebrated his ninetieth birthday. Moreover, the year 2013 is also the seventieth anniversary of the publication of his first scientific paper. For the past decade, Jaroslav has been spending his retirement in fruitful work on his beloved , resulting in 38 scientific papers devoted to the taxonomy and chorology of that group. In acknowledgement of Jaroslav’s life achievements in entomology, he was appointed a honorary member of the Czech Entomological Society in 2006. We would like to congratulate Jaroslav Stehlík on his anniversaries and wish him all the best for the years to come. Here we include some recollections by two of his close colleagues, to commemorate the occasion.

Petr Kment

Meeting Jaroslav Stehlík Two research topics have dominated the scientific life of Jaroslav Stehlík. One is eco-faunistic research into the Heteroptera of Moravia and Slovakia, which has resulted in a plethora of publications containing exemplary summaries of the chorology, bionomics and diagnostics of concerned. The other has been the alpha-taxonomy of the Pyrrhocoridae (firebugs, red bugs, cotton-stainers) and Largidae of the world. Jaroslav was also able to maintain an overview of general entomological approaches to the true bugs, although he did not actively pursuit such studies. I cannot really remember how I perceived Jaroslav’s abilities when, in early 1950s I was casting about for a suitable topic for my MSc thesis (“diploma thesis” in those days) at the Biological Faculty of Charles University in Prague. From my point of view, it should have been on Heteroptera but could not address taxonomy (a subject then very nearly taboo to the authorities). I was not interested in diapause (then a favoured subject) and I flatly refused applied topics. I asked Jaroslav for advice, warmly embraced his suggestion, and in 1957 I defended a thesis entitled “Ectodermal female genitalia of the trichophorous Heteroptera”. Evidence of the topicality of the subject was that, almost simultaneously, two other university students of the Heteroptera, Geoffrey G. E. Scudder in Oxford and Carl W. Schaefer in Storrs, Connecticut, began their theses along similar lines. I cannot be grateful enough to Jaroslav for setting me on my life-time research course, a combination of morphology, phylogeny, higher classification and taxonomy of critical groups of true bugs. My initial contacts with Jaroslav could, of course, never be entirely free of the limits imposed by our differences in age and status, but these soon disappeared and I had the privilege to know him both as a heteropterist and a personality. He was always true to his

5 P. K MENT, P. ŠTYS, H. GÜNTHER pronounced opinions on science, but he did not expect you to share them; he was friendly and open with those he trusted, but always ready to defend his territory. He loved the research he had chosen for himself, loved the Moravian and Slovakian countryside and its natural history, and in times when boorishness was nearly a required norm, he remained courteous and elegant, a trait surely greatly appreciated by his women collaborators. He did not cease research on retirement, and the volume of flow of his publications on Pyrrhocoroidea is astonishing. Surely all hemipterists would appreciate Jaroslav’s decision to write a kind of synthesis of his unique knowledge of these bugs. Dear Jaroslav, I wish you health, strength and happiness in both your personal and scientific life, and many further years of enjoyment of your research.

Pavel Štys

A circuit tour and short circuits in southern Slovakia with Jaroslav in 1991 At the invitation of my friend Jaroslav Stehlík we undertook a collecting trip through South Slovakia in June 1991. It was a highly interesting and successful trip with plenty of interesting collection sites and rare species of bugs to capture. Starting in Brno, we first went to the Nové Zámky district with all the famous places in its surroundings. We prepared breakfast in our hotel room and for this purpose Jaroslav had an immersion heater in his luggage. Trying to heat about a litre of water to prepare the coffee, there was a short in the unit and we had coffee neither that morning nor for nearly all the days that followed. Jaroslav was quite upset; the immersion heater belonged his wife, who used it in the school in which she taught. It was not possible to obtain a replacement. This was our first mishap with the electricity. We drove on and took up residence in a hotel in Ve¾ký Krtíš. It was very hot, even during the night, and we kept the window open. This type of window could rotate about its vertical axis. In the night, a strong wind blew up, the window spun and threw the bedside lamp to the ground. In an instant, the whole hotel had no power. A classic case of short circuit. Our last hotel was located in Luèenec, farther to the east. At that time Luèenec was a pretty, small town with many shops and I decided to look for an immersion heater to replace Madame Stehlík’s ruined one. I was successful, and managed to buy an identical model in a hardware shop. An hour later Jaroslav came back to the hotel, again very sad because he had not obtained an immersion heater in the very same shop. The shopkeeper had informed him that the article he required had been sold only a short time earlier and that it had been the only one they had. But then his joy and mine were unconfined, when I was able to tell him who the previous customer had been, and hand the device to him. Back home in Ingelheim, mounting and identifying the bugs collected for the first time, my eyes were opened to the species, most of which were new for my collection. A list of plant-bug species new for Slovakia was published in Entomological Problems (GÜNTHER 2000). We collected many more rare species during those days in June 1991, a proof that even electrical short circuits could not influence our success.

Hannes Günther

6 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Jaroslav L. Stehlík

On a collecting excursion with Slovak heteropterists, 1960s. From left to right: J. Vacek, Irina Vavøínová, Adela Beláková, Jaroslav L. Stehlík, and Olga Štìpanovièová (archive of the Moravian Museum, Brno).

The staff of the Dept. Entomology, Moravian Museum on the roof of its building in 1969. From left to right: Pavel Lauterer, Marie Novotná, Ludmila Pospíšilová, Jaroslav L. Stehlík, Irina Vavøínová, and Mrs. Kaczmarová (archive of the Moravian Museum, Brno).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 7 P. K MENT, P. ŠTYS, H. GÜNTHER

Jaroslav L. Stehlík in his office in the Moravian Museum in early 1950s (archive of the Moravian Museum, Brno).

Bibliography (2003–2013) For a complete bibliography of Jaroslav L. Stehlík in the period 1943–2002 see ŠTEPANOVIÈOVÁ & BIANCHI (2003).

STEHLÍK J. L. 2003: Largidae and Pyrrhocoridae of Nepal (Heteroptera). Entomologica Basiliensia 25: 1–11. STEHLÍK J. L. & JINDRA Z. 2003: Largidae and Pyrrhocoridae of Thailand. Acta Musei Moraviae, Scientiae Biologicae 88: 5–19. STEHLÍK J. L. 2005a: Largidae and Pyrrhocoridae of Laos (Hemiptera: Heteroptera). Folia Heyrovskyana 12 (2004): 141–160. STEHLÍK J. L. 2005b: Largidae and Pyrrhocoridae collected by Alexander Riedel in Irian Jaya (New Guinea) from 1990 up to 1996 (Heteroptera). Linzer Biologische Beiträge 37: 1719–1736.

8 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Jaroslav L. Stehlík

STEHLÍK J. L. 2006a: New taxa of Pyrrhocoroidea (Heteroptera) from the Oriental Region in the Natural History Museum in London. Pp. 653–680. In: RABITSCH W. (ed.): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia 19: 1–1184. STEHLÍK J. L. 2006b: Four new species of the Neodindymus Stehlík, 1965 (Heteroptera, Pyrrhocoridae). Entomologica Basiliensia et Collectionis Frey 28: 33–48. STEHLÍK J. L. 2006c: Four new species of the Neotropical genus Theraneis Spinola, 1837 (Heteroptera: Largidae, Larginae). Entomologica Basiliensia et Collectionis Frey 28: 49–58. STEHLÍK J. L. & JINDRA Z. 2006a: Five new species of the genus Dindymus (Heteroptera: Pyrrhocoridae). Acta Entomologica Musei Nationalis Pragae 46: 21–30. STEHLÍK J. L. & JINDRA Z. 2006b: New species of Largidae and Pyrrhocoridae (Heteroptera) from the Oriental region. Acta Entomologica Musei Nationalis Pragae 46: 31–41. STEHLÍK J. L. & JINDRA Z. 2006c: Notes on some Oriental species of the genus Dysdercus Guérin Méneville, 1831 (Pyrrhocoridae, Heteroptera). Acta Musei Moraviae, Scientiae Biologicae 91: 53–60. STEHLÍK J. L. & JINDRA Z. 2006d: A revision of the genus Ascopocoris Stehlík et Kerzhner, 1999 (Pyrrhocoridae, Heteroptera). Acta Musei Moraviae, Scientiae Biologicae 91: 61–68. STEHLÍK J. L. 2007a: On the Afrotropical species of Myrmoplasta (Heteroptera: Pyrrhocoridae). Acta Entomologica Musei Nationalis Pragae 47: 93–108. STEHLÍK J. L. 2007b: On the Malagasy genus Jourdainana Distant, 1913 (Heteroptera, Pyrrhocoridae). Entomologica Basiliensia et Collectionis Frey 29: 1–12. STEHLÍK J. L. 2007c: Brancucciana Ahmad et Zaidi, 1986 is the valid name for Ascopocoris Stehlík et Kerzhner, 1999 (Heteroptera: Pyrrhocoridae). Acta Musei Moraviae, Scientiae Biologicae 92: 109–110. STEHLÍK J. L. 2007d: New combinations in the genus Rosaphe Kirkaldy et Edwards, 1902 (= Astemma auct., nec Le Peletier et Serville, 1825) (Heteroptera: Largidae, Larginae). Acta Musei Moraviae, Scientiae Biologicae 92: 111–113. STEHLÍK J. L. 2007e: Largidae and Pyrrhocoridae (Heteroptera) of Meghalaya state, India. Acta Musei Moraviae, Scientiae Biologicae 92: 115–129. STEHLÍK J. L. 2007f: Two new species and new records of Pyrrhocoroidea from Laos (Heteroptera). Acta Musei Moraviae, Scientiae Biologicae 92: 131–136. STEHLÍK J. L. & JINDRA Z. 2007a: Five new species of the genus Dindymus Stål from New Guinea (Pyrrhocoridae, Heteroptera). Pp. 145–152. In: RENKER C. (ed.): Festschrift zum 70. Geburtstag von Hannes Günther. Mainzer Naturwissenschaftliche Archiv, Beiheft 31: 1–339. STEHLÍK J. L. & JINDRA Z. 2007b: Largulini – a new tribe of Larginae from Jamaica (Heteroptera, Largidae). Entomologica Basiliensia et Collectionis Frey 29: 13–20. STEHLÍK J. L. 2008: New taxa of Afrotropical Pyrrhocoridae (Hemiptera: Heteroptera). Entomologica Basiliensia et Collectionis Frey 30: 3–20. STEHLÍK J. L. & JINDRA Z. 2008a: New taxa of the Largidae and Pyrrhocoridae (Hemiptera: Heteroptera) from the Oriental Region. Acta Entomologica Musei Nationalis Pragae 48: 611–648. STEHLÍK J. L. & JINDRA Z. 2008b: A contribution to knowledge of Pyrrhocoroidea (Heteroptera) of Zambia. Acta Musei Moraviae, Scientiae Biologicae 93: 29–42. STEHLÍK J. L. & KMENT P. 2008: Myrmoplastoides subgen. nov. of the genus Myrmoplasta (Hemiptera: Heteroptera: Pyrrhocoridae) from the Oriental region. Zootaxa 1782: 61–64. STEHLÍK J. L. 2009a: Pseudodindymus, a new subgenus of Dindymus (Hemiptera: Heteroptera: Pyrrhocoridae) from Oriental Region. Zootaxa 2010: 1–15. STEHLÍK J. L. 2009b: Five new species of Antilochus Stål, 1863 from the Old World (Heteroptera: Pyrrhocoridae). Acta Musei Moraviae, Scientiae Biologicae 94: 57–71. STEHLÍK J. L. 2009c: Two new species of the genus Sericocoris Karsch, 1892 from Central Africa (Hemiptera, Heteroptera, Pyrrhocoridae). Entomologica Basiliensia et Collectionis Frey 31: 1–9. STEHLÍK J. L. & JINDRA Z. 2010a: Two new species of the genus Roscius (Hemiptera: Heteroptera: Pyrrhocoridae). Zootaxa 2651: 59–63. STEHLÍK J. L. & JINDRA Z. 2010b: Two new species of the genus Sericocoris (Hemiptera: Heteroptera: Pyrrhocoridae) from Cameroon. Acta Musei Moraviae, Scientiae Biologicae 95(2): 19–23. STEHLÍK J. L. & KMENT P. 2010: Largus giganteus sp. nov. from Brazil and notes on hybridization within Largus (Hemiptera: Heteroptera: Largidae). Acta Entomologica Musei Nationalis Pragae 50: 53–58.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 9 P. K MENT, P. ŠTYS, H. GÜNTHER

STEHLÍK J. L. & BRAILOVSKY H. 2011: Two new genera of the tribe Largulini (Hemiptera: Heteroptera: Largidae) from Greater Antilles. Acta Entomologica Musei Nationalis Pragae 51: 449–456. STEHLÍK J. L. & JINDRA Z. 2011: Schaeferiana (Gaboniella subgen. n.) incompleta sp. n. from Gabon, with notes on its relationships and new records from the Central African Republic (Hemiptera, Heteroptera, Pyrrhocoridae). ZooKeys 126: 49–56. STEHLÍK J. L. & KMENT P. 2011a: Antilochus (Neaeretus) pterobrachys sp. nov. and the correct name of the subgenus Afroantilochus (Hemiptera: Heteroptera: Pyrrhocoridae). Acta Entomologica Musei Nationalis Pragae 51: 49–53. STEHLÍK J. L. & KMENT P. 2011b: Redescription of Pararhaphe and review of Arhaphe (Hemiptera: Heteroptera: Largidae) of America north of Mexico. Zootaxa 3058: 35–54. STEHLÍK J. L. & JINDRA Z. 2012: Australodindymus nigroruber gen. et sp. nov. from Western Australia (Hemiptera: Heteroptera: Pyrrhocoridae). Zootaxa 3316: 57–62. STEHLÍK J. L. & KMENT P. 2012: Description of Physopelta finisterrae sp. nov. from Papua New Guinea (Hemiptera: Heteroptera: Largidae). Acta Entomologica Musei Nationalis Pragae 52: 349–354. SCHAEFER C. W. & STEHLÍK J. L. 2013: Caribbean Sea Region Pyrrhocoroidea (Hemiptera: Pyrrhocoridae, Largidae). Neotropical Entomology 42: 372–383. STEHLÍK J. L. 2013a: A description of four new species of Dindymus (Hemiptera: Heteroptera: Pyrrhocoridae) from south-eastern Asia. Acta Musei Moraviae, Scientiae Biologicae 98(1): 41–51. STEHLÍK J. L. 2013b: Review and reclassification of the Old World genus Physopelta (Hemiptera: Heteroptera: Largidae). Acta Entomologica Musei Nationalis Pragae 53: 505–584.

Petr Kment

References

GÜNTHER H. 2000: Contribution to the fauna of plant bugs (Heteroptera: ) in Slovakia. Entomological Problems 31: 59–60. ŠTEPANOVIÈOVÁ O. & BIANCHI Z. 2003: K 80. narozeninám RNDr. Jaroslava L. Stehlíka, CSc. (On the occasion of the 80th birthday of RNDr. Jaroslav L. Stehlík, CSc. Klapalekiana 39: 189–197.

10 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 11–19, 2013

RNDr. Pavel Lauterer, on the occasion of his eightieth birthday

On 25 August 2013, RNDr. Pavel Lauterer celebrated a remarkable anniversary. His eighty years have been marked by hundreds of fundamental scientific contributions, particularly (but not only) on jumping plant-lice, leafhoppers, and on their natural enemies; by hundreds of thousands of mounted and identified specimens in collections; and by the hundreds of students he inspired into entomology through his indefatigable enthusiasm for insects and nature in general, his sense of humour and his exceptional friendly and encouraging attitude. As one of the most influential personalities in the field, he was appointed Honorary Member of the Czech Entomological Society in 2006. International recognition for his qualities is inseparable from the contributions in this Festschrift. For many of us who know him personally, Pavel is a respected colleague, an excellent teacher and a good friend. We wish him a very happy birthday.

Igor Malenovský

It was in April 1981 that I had the privilege of meeting Pavel the first time personally, although we had already maintained a correspondence for some three years. At that time I was working on my Ph.D thesis on psyllids and had reached a point at which questions had accumulated to the point at which expert input was becoming essential. I was very grateful that Pavel could extend his help and that I could visit him in Brno. From Brno railway station, where he kindly picked me up, we went straight to the Entomology Department of the Moravian Museum, situated in a former residential house, and started looking at specimens and talking about psyllids. I was impressed by the huge psyllid collection he had assembled over the years and even more by his profound knowledge of the biology, biogeography and taxonomy of the psyllids, and of plants and in general. Pavel also showed me his rearing experiments, confined to the minute balcony of his home apartment. Even though the setting

Pavel Lauterer collecting insects in the Belianské Tatry Mts. in Slovakia in 1962 (photo Mrs. Vìra Lautererová).

11 I. MALENOVSKÝ, D. BURCKHARDT, S. DROSOPOULOS

was simple, he made many invaluable observations and shed light on some very vexed issues, such as the identities of Heterotrioza horvathii and obliqua as forms of H. chenopodii arising out of different photoperiods. I also remember the wonderful Czech meals and excellent beer we had after long days of work. This visit marked the start of a highly fruitful Pavel Lauterer and Reinhard Remane collecting leafhoppers on the collaboration that has lasted shore of the Meugliano Lake during the 14th Central European to this very day, with the Auchenorrhyncha Meeting in Ivrea, , 2007 (photo Gernot Kunz). description of a new Trioza species which, independently, we both named Trioza laserpitii. Pavel is a true field entomologist and one of the best collectors I know. Thanks to his fantastic knowledge of biology, he finds the rarest and most sought-after insects. One would not be guilty of exaggeration to observe that if Pavel does not discover a particular species in a certain locality, the chances are very high that it does not occur there. I remember with great pleasure our highly successful joint excursions to the xerothermic rock face of Les Follatères in the Lower Valais of , and to the species-rich subalpine biotopes on Rigi and Pilatus in the pre-Alps of central Switzerland. In the evenings of any excursion Pavel would take hours to process the large quantities of specimens that he had collected during the day. It is no surprise that these collections contained several new psyllid species, which we described together in the following years. In fact, many of our joint projects started with a new species which needed description and then developed into a genuine revision of the entire group: our papers on the , and the exilis group are examples. Pavel, I would like to thank you here, with all my heart, for your generous help, for continuous incentives and for the friendship we have shared over the years.

Daniel Burckhardt

It is with pleasure that I think back to the time when Pavel first visited Greece, nearly 20 years ago, to the collecting trips we took together, and to his infectious enthusiasm. Visiting the area of Lake Mornos, close to my own village of Skaloula, he exclaimed: “Sakis, I have never in my life seen better biotopes!” Lake Mornos is a storage reservoir, supplying water to the city of Athens. The surrounding area is sparsely populated and undisturbed and I have submitted a proposal to turn it into a National Park.

12 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Pavel Lauterer

Irina Vavøínová (left), Jaroslav L. Stehlík and Pavel Lauterer hosting specialist David A. Young (right) on his visit of the Moravian Museum in 1962.

Further, even when he was staying in the city of Athens, he would waste no opportunity to collect. He did so on the premises of the Benaki Phytopathological Institute, in my garden in Kiphisia and in the Botanical gardens in the centre of Athens, where he once stayed on, still collecting, right up to sunset. Stopping when the light failed, he found himself locked in as the gates had been closed long before, and a guard had to be called to set him free. Of course, we have collected new Hemiptera species and described them together, but quite apart from the scientific importance of his visits I also remember him for his wonderful sense of humour. On a visit to the Peloponnese, after a day of hard work collecting on Mt. Parnon we were relaxing by the seaside in Paralion Astros, enjoying ouzo with fine cooked cephalopod and I asked him if it was the first time he had eaten octopus. His answer was negative; it was in fact the second time. So I asked him where then he had had octopus before and he answered: “At home, from a tin.” So I said: “Pavel, then now you eat octopus for the first time” and of course ordered more. I also remember how he used to be pleased with the abundance of fresh lemon served with a meal, as lemons used to be very scarce in his country. It hardly needs saying that Pavel is a person I will never forget.

Sakis Drosopoulos

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 13 I. MALENOVSKÝ, D. BURCKHARDT, S. DROSOPOULOS

Bibliography (2003–2013) For a complete bibliography of Pavel Lauterer in the period 1954–2002 see MALENOVSKÝ & BURCKHARDT (2003): Pavel Lauterer at Seventy. Acta Musei Moraviae, Scientiae biologicae 88: 181–190.

120. BURCKHARDT D. & LAUTERER P. 2003: Verzeichnis der Blattflöhe (Psylloidea) Deutschlands. Pp. 155–164. In: KLAUSNITZER B. (ed.): Entomofauna Germanica Band 6. Entomologische Nachrichten und Berichte Suppl. 8: 1–344. 121. LAUTERER P. 2003a: First reports of some Pipunculidae (Diptera) from the Czech Republic and Slovakia with a note from . Acta Musei Moraviae, Scientiae biologicae 88: 31–35. 122. LAUTERER P. 2003b: Poznámky o nìkterých druzích rodu Formica na hranicích okresù Blansko a Zïár nad Sázavou (Èeská republika). (Notes of several findings of of the genus Formica in frontiers of the districts Blansko and Žïár nad Sázavou.) Formica 6: 54–59 (in Czech, English summary). 123. LAUTERER P. 2003c: Zachraòme mravence microcephalum na jižní Moravì. (Save the species Liometopum microcephalum in southern Moravia.) Formica 6: 69–71 (in Czech, English summary). 124. NICKEL H., BILLEN W., GÜNTHART H., LAUTERER P., LOCKER H., MALENOVSKÝ I., MÜHLETHALER R., SCHÜRRER B. & WITSACK W. 2003: Zur Fauna der Zikaden, Wanzen und Augenfliegen des Kaiserstuhls (Hemiptera: Auchenorrhyncha et Heteroptera; Diptera: Pipunculidae). Beiträge zur Zikadenkunde 6: 39–46. 125. NICKEL H., HOLZINGER W., LAUTERER P., REMANE R. & WITSACK W. 2003: Die Spornzikaden-Gattung Metropis Fieber, 1866 in Mitteleuropa (Hemiptera: Fulgoromorpha: ). Beiträge zur Zikadenkunde 6: 47–52. 126. PREISLER J. & LAUTERER P. 2003: Some new species of planthoppers and leafhoppers for the Czech Republic and Slovakia (Hemiptera, Auchenorrhyncha). Beiträge zur Zikadenkunde 6: 53–56. 127. WYNIGER D., MÜHLETHALER R., LAUTERER P. & BURCKHARDT D. 2003: Blattflöhe, Zikaden und Landwanzen (Hemiptera) im Naturschutzgebiet Wildenstein. Mitteilungen der Naturforschenden Gesellschaften beider Basel 7: 293–304. 128. FIALOVÁ R., NAVRÁTIL M., VÁLOVÁ P., LAUTERER P., K OCOUREK F. & PONCAROVÁ-VORÁÈKOVÁ Z. 2004: Epidemiology of European stone fruit yellows in the Czech Republic. Acta Horticulturae 657: 483–487. 129. GÜNTHART H., MÜHLETHALER R. & LAUTERER P. 2004: Für die Schweiz neue Zikadenarten und Ergänzungen zu bereits bekannten Arten (Hemiptera, Auchenorrhyncha). Mitteilungen der Entomologischen Gesellschaft Basel 54: 150–160. 130. LAUTERER P. 2004a: Subfosilie èlenovcù a žížal v Èeské republice. [Subfossil and earthworms in the Czech Republic.] Pp. 86–87. In: BRYJA J. & ZUKAL J. (eds.): Zoologické dny 2004. Sborník abstraktù z konference 12.–13. února 2004. Ústav biologie obratlovcù AV ÈR, Brno, 232 pp (in Czech). 131. LAUTERER P. 2004b: Eneolitické a støedovìké nálezy mravencù (, Formicidae) v Èeské republice. (Eneolitic and medieval ant findings in Czech republic.) Formica 7: 71–75 (in Czech, English summary). 132. LAUTERER P. 2004c: Køísi – Auchenorrhyncha. Pp. 75–76. In: JÓŽA M. & VONIÈKA P. et al.: Jizerskohorská rašeliništì. [Peatbogs of the Jizerské hory Mts.] Jizersko-ještìdský horský spolek, Liberec, 159 pp, 1 map (in Czech). 133. LAUTERER P. 2004d: Nejvýznaènìjší druhy hmyzu vrchu Kvìtnice. [The most noteworthy insect species of the Kvìtnice Hill.] Sborník Muzea Brnìnska 2004: 154–162 (in Czech). 134. LAUTERER P. 2004e: Vzpomínky na Dr. Ilju Okáliho, Csc. [Remembering Dr. Ilja Okáli.] Pp. 27–33. In: HOLÈÍK J. (ed.): Ilja Okáli. Edicia medailony. Múzeum, Metodický, študijný a informaèný èasopis pre pracovníkov múzeí a galérií Suppl. 2004: 1–44 pp. 135. LAUTERER P. & BURCKHARDT D. 2004: The West Palaearctic species of the flavipennis (Foerster) complex (Hemiptera, Psylloidea). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 77: 261–275.

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136. NAVRÁTIL M., FIALOVÁ R., KOCOUREK F., LAUTERER P., VÁLOVÁ P., ŠAFÁØOVÁ D. & PONCAROVÁ- VORÁÈKOVÁ Z. 2004: Problems on European stone fruit yellows in the Czech Republic. Acta Fytotechnica et Zootechnica (Nitra) 7: 217–219. 137. BATELKA J., STRAKA J., MALENOVSKÝ I. & LAUTERER P. 2005: (øasnokøídlí). Pp. 559–561. In: FARKAÈ J., KRÁL D. & ŠKORPÍK M. (eds.): Èervený seznam ohrožených druhù Èeské republiky. Bezobratlí. Red list of threatened species in the Czech Republic. Invertebrates. Agentura ochrany pøírody ÈR, Praha, 760 pp (in Czech and English). 138. KOCOUREK F. & LAUTERER P. 2005: QD 1048. Systémy integrované ochrany ovocných døevin proti komplexu škodlivých organismù. Metodika pro monitoring mery pruni jako pøenašeèe ESFY a podklady pro ochranu vùèi pøenašeèi ESFY. [Methods for monitoring of the psyllid Cacopsylla pruni as a vector of ESFY and data for protection against the vector of ESFY.] VÚRV, Praha-Ruzynì, 9 pp (in Czech). Available online: http://www.vurv.cz/files/Publications/Metodika_Cpruni.pdf 139. LAUTERER P. 2005: New records and hosts for the parasitoid Halictophagus agalliae (Strepsiptera: ). Acta Musei Moraviae, Scientiae biologicae 90: 215–217. 140. LAUTERER P. & M ALENOVSKÝ I. 2005: Psylloidea (mery). Pp. 156–159. In: FARKAÈ J., KRÁL D. & ŠKORPÍK M. (eds.): Èervený seznam ohrožených druhù Èeské republiky. Bezobratlí. Red list of threatened species in the Czech Republic. Invertebrates. Agentura ochrany pøírody ÈR, Praha, 760 pp (in Czech and English). 141. MALENOVSKÝ I. & LAUTERER P. 2005a: Auchenorrhyncha (køísi). Pp. 147–155. In: FARKAÈ J., KRÁL D. & ŠKORPÍK M. (eds.): Èervený seznam ohrožených druhù Èeské republiky. Bezobratlí. Red list of threatened species in the Czech Republic. Invertebrates. Agentura ochrany pøírody ÈR, Praha, 760 pp (in Czech and English). 142. MALENOVSKÝ I. & LAUTERER P. 2005b: Leafhoppers, planthoppers and psyllids (Hemiptera, , Fulgoromorpha, Psylloidea) in ruderal habitats: material attracted by light in the suburbs of Brno (Czech Republic). Acta Musei Moraviae, Scientiae biologicae 90: 195–207. 143. BØEZÍKOVÁ M. & LAUTERER P. 2006a: Žilnatka vironosná (Hyalesthes obsoletus Signoret, 1865) (Hemiptera: ), významný pøenašeè fytoplamy stolburu bramboru. [The Hyalesthes obsoletus Signoret, 1865 (Hemiptera: Cixiidae), a significant vector of the potato stolbur phytoplasma.] Agro 2006(7): 22–24 (in Czech). 144. BØEZÍKOVÁ M. & LAUTERER P. 2006b: Žilnatka vironosná (Hyalesthes obsoletus Signoret, 1865) (Fulgoromorpha: Cixiidae) – rozšíøení, biologie a význam pro pøenos fytoplasmy stolburu bramboru. The planthopper Hyalesthes obsoletus Signoret, 1865 (Fulgoromorpha, Cixiidae) – spread, biology and importance for transmission of Potato stolbur phytoplasma. Pp. 91–92. In: XVII. Èeská a slovenská konference o ochranì rostlin: sborník abstraktù. 12.–14. záøí 2006, Praha. XVII. Czech and Slovak Plant Protection Conference: book of abstracts. 12–14th September 2006, Prague. Èeská zemìdìlská univerzita, Praha, 386 + 4 + 20 pp + CD-ROM (in Czech and English). 145. BURCKHARDT D. & LAUTERER P. 2006: The Palaearctic Triozids associated with Rubiaceae (Hemiptera, Psylloidea): a taxonomic re-evaluation of the Trioza galii Foerster complex. Revue Suisse de Zoologie 113: 269–286. 146. ÈERMÁK V. & LAUTERER P. 2006: Pøíspìvek k bionomii mer Cacopsylla picta a . [Contribution to the bionomics of the psyllids Cacopsylla picta and Cacopsylla melanoneura.] P. 75. In: BRYJA J. & ZUKAL J. (eds.): Zoologické dny Brno 2006. Sborník abstraktù z konference 9.–10. února 2006. Ústav biologie obratlovcù AV ÈR, Brno, 268 pp (in Czech). 147. FIALOVÁ R., NAVRÁTIL M., LAUTERER P. & V ÁLOVÁ P. 2006: Molekulární diagnostika fytoplazmy proliferace jabloní a hmyzích vektorù v jabloòových sadech Èeské republiky. Molecular tests to determine apple proliferation phytoplasma presence in psyllid vectors from apple orchards in the Czech Republic. P. 164. In: XVII. Èeská a slovenská konference o ochranì rostlin: sborník abstraktù. 12.–14. záøí 2006, Praha. XVII. Czech and Slovak Plant Protection Conference: book of abstracts. 12–14th September 2006, Prague. Èeská zemìdìlská univerzita, Praha, 386 + 4 + 20 pp + CD-ROM (in Czech and English). 148. LAUTERER P. 2006a: Možnosti využití sacích pastí Státní rostlinolékaøské správy pro kontrolu a signalisaci mšic pro sledováni rùzných skupin èlenovcù. [Possibilities of use of suction traps of the State phytosanitary service for control and signalisation of for monitoring of various groups of arthropods.]. Pp. 104–105. In: BRYJA J. & ZUKAL J. (eds.): Zoologické dny Brno 2006. Sborník abstraktù z konference 9.–10. února 2006. Ústav biologie obratlovcù AV ÈR, Brno, 268 pp (in Czech).

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149. LAUTERER P. 2006b: Pipunculidae Walker, 1834. In: JEDLIÈKA L., STLOUKALOVÁ V., KÚDELA M. (eds.): Checklist of Diptera of the Czech Republic and Slovakia. Electronic version 1. Available online: http://zoology.fns.uniba.sk/diptera2006 and on CD-ROM (ISBN 80-969629-0-6). 150. LAUTERER P. & B ØEZÍKOVÁ M. 2006: Nové šíøení køísa Hyalesthes obsoletus Signoret, 1865 (Fulgoromorpha: Cixiidae) v Èeské republice. [New spreading of the planthopper Hyalesthes obsoletus Signoret, 1865 (Fulgoromorpha: Cixiidae) in the Czech Republic.] P. 105. In: BRYJA J. & ZUKAL J. (eds.): Zoologické dny Brno 2006. Sborník abstraktù z konference 9.–10. února 2006. Ústav biologie obratlovcù AV ÈR, Brno, 268 pp (in Czech). 151. LAUTERER P. & ÈERMÁK V. 2006a: Mery – vektory fytoplazem clusteru AP. Psyllids – the vectors of the AP cluster . Pp. 100–101. In: XVII. Èeská a slovenská konference o ochranì rostlin: sborník abstraktù. 12.–14. záøí 2006, Praha. XVII. Czech and Slovak Plant Protection Conference: book of abstracts. 12–14th September 2006, Prague. Èeská zemìdìlská univerzita, Praha, 386 + 4 + 20 pp + CD- ROM (in Czech and English). 152. LAUTERER P. & ÈERMÁK V. 2006b: Biology of the psyllid vectors of AP and ESFY phytoplasmas (Hemiptera, Psylloidea). Pp. 482–487. In: Sborník pøíspìvkù. XVII. èeská a slovenská konference o ochranì rostlin. 12. –14. záøí Praha 2006. Proceedings XVII. Czech and Slovak Plant Protection Conference. Èeská zemìdìlská univerzita v Praze, Katedra ochrany rostlin, 640 pp. 153. LAUTERER P. & M ALENOVSKÝ I. 2006a: Hlavatìnkovití (Pipunculidae, Diptera) CHKO Kokoøínsko. (Big- headed (Pipunculidae, Diptera) of Kokoøínsko Protected Landscpe Area.) Bohemia centralis 27: 457–466 (in Czech, English abstract). 154. LAUTERER P. & M ALENOVSKÝ I. 2006b: New records of Pipunculidae (Diptera) from the Czech Republic. Entomofauna Carpathica 18: 37–38. 155. LAUTERER P. & M ALENOVSKÝ I. 2006c: hyalinus (Osborn, 1900) – køísek. Pp. 267–268. pruinosa Say, 1830 – voskovka zavleèená. Pp. 268–268. buxi (Linnaeus, 1758) – mera zimostrázová. P. 289. In: MLÍKOVSKÝ J. & STÝBLO P. (eds.): Nepùvodní druhy fauny a flóry Èeské republiky. [Alien species in the fauna and flora of the Czech Republic.] Èeský svaz ochráncù pøírody, Praha, 496 pp. 156. MALENOVSKÝ I. & LAUTERER P. 2006: Cicadomorpha – køísci a Fulgoromorpha – svítilky. Pp. 266–267. Stictocephala bisonia Kopp & Yonke, 1977 – ostnohøbetka ovocná. Pp. 269–271. melissae Curtis, 1837 – pidikøísek. P. 281. Psylloidea – Mery. P. 274. Arytaina genistae (Latreille, 1804) – mera janovcová. P. 277. Cacopsylla hippophaes (Förster, 1848) – mera rakytníková. Pp. 278–279. Livilla variegata (Löw, 1881) – mera štìdøencová. Pp. 285–286. In: MLÍKOVSKÝ J. & STÝBLO P. (eds.): Nepùvodní druhy fauny a flóry Èeské republiky. [Alien species in the fauna and flora of the Czech Republic.] Èeský svaz ochráncù pøírody, Praha, 496 pp. 157. BURCKHARDT D., LAUTERER P. & NIEDRINGHAUS R. 2007: Die Blattflöhe der Sammlung Struve von der Nordseeinsel Borkum (Hemiptera, Psylloidea). Beiträge zur Zikadenkunde 9: 39–44. 158. FIALOVÁ R., NAVRÁTIL M., LAUTERER P. & NAVRKALOVÁ V. 2007: “Candidatus Phytoplasma prunorum”, the phytoplasma infection of Cacopsylla pruni from orchards and from overwintering habitats in Moravia (Czech Republic). Bulletin of Insectology 60: 183–184. 159. KUÈEROVÁ J., TALÁCKO L., LAUTERER P., NAVRÁTIL M. & FIALOVÁ R. 2007: Molecular tests to determine “Candidatus Phytoplasma pyri” presence in psyllid vectors from a pear tree orchard in the Czech Republic – a preliminary report. Bulletin of Insectology 60: 191–192. 160. LAUTERER P. 2007a: Vzpoura otrokù - mravenèí Spartakové. (Sklavenausfand – Spartakus unter den Ameisen?). Formica 10: 19–22 (in Czech, German summary). 161. LAUTERER P. 2007b: Mravenci Lasius niger (Linné) a larvy mšicožravých pestøenek (Diptera, Syrphidae). (Ameisen Lasius niger (L.), Blattläuse und Larven der Blattläusefressenden Schwebfliegen (Diptera, Syrphidae)). Formica 10: 29–32 (in Czech, German summary). 162. LAUTERER P. & ÈERMÁK V. 2007: Overwintering of psyllids (Hemiptera, Psylloidea) – the vectors of the phytoplasma apple proliferation cluster. P. 76. In: BRYJA J., ZUKAL J & ØEHÁK Z. (eds.): Zoologické dny 2007. Sborník abstraktù z konference 8.–9. února 2007. Ústav biologie obratlovcù AV ÈR, Brno, 224 pp. 163. LAUTERER P. & DOROW W. H. O. 2007: 3.6 Psylloidea (Blattflöhe). In: DORROW W. H. O. & KOEPOLKE J. P. (eds.): Hohestein. Zoologische Untersuchungen 1994–1996, Teil 2. Mitteilungen der Hessischen Landesforstverwaltung 42: 95–101.

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164. Mühlethaler R., Günthart H., Burckhardt D., Holzinger W., Kunz G., Lauterer P., Maczey N., Malenovský I., Nickel H., Niedringhaus R., Seljak G., Walter S. & Witsack W. 2007: Zur Fauna der Zikaden, Blattfl6he und deren Parasitoide der Alp Flix (Graubünden, Schweiz). (Hemiptera: Auchenorrhyncha et Psylloidea; Diptera: Pipunculidae, Strepsiptera: Elenchidae). Beiträge zur Zikadenkunde 9: 45–53. 165. SUCHÁ J., TALÁCKO L., LAUTERER P., NAVRÁTIL M. & FIALOVÁ R. 2007: Molekulární determinace pøítomnosti pear decline fytoplazem v merách vyskytujících se ve výsadbì hrušní v Èeské republice – pøedbìžná zpráva. [Molecular determination of a presence of pear decline phytoplasmas in psyllids occurring in pear tree plantations in the Czech Republic – a preliminary report.] Pp. 53–58. In: Inovace pìstování ovocných plodin. [Inovation of fruit growing.] Výzkumný a šlechtitelský ústav ovocnáøský Holovousy, s.r.o., Holovous, 214 pp (in Czech). 166. BURCKHARDT D., LAUTERER P. & NIEDRINGHAUS R. 2008: Die Pflanzenläuse der Ostfriesischen Inseln (Hemiptera: ). Zum Kenntnisstand über die Fauna der Blattflöhe, Schild- und Blattläuse (Psylloidea, Coccoidea, Aphidoidea). Schriftenreihe Nationalpark Niedersächsische Wattenmeer 11: 179–184. 167. BURCKHARDT D., MÜHLETHALER R., LAUTERER P., MALENOVSKÝ I. & KUNZ G. (2008): Die Blattflöhe und Zikaden der Alp Flix (Insecta, Hemiptera: Psylloidea und Auchenorrhyncha). Jahresbericht der Naturforschenden Gesellschaft Graubünden 114(2007): 91–97. 168. ÈERMÁK V. & LAUTERER P. 2008: Overwintering of psyllids in South Moravia (Czech Republic) with respect of the vectors of the Apple Proliferation cluster phytoplasmas. Bulletin of Insectology 61: 147–148. 169. FIALOVÁ R., NAVRÁTIL M., VÁLOVÁ P. & LAUTERER P. 2008: Molecular tests to determine Apple Proliferation Phytoplasma presence in psyllid vectors from apple tree orchards in the Czech Republic. In: CAGLAYAN K. & ERTUNC F. (eds.): Proceedings of the XXth International Symposium on - and Virus- like Diseases of Temperate Fruit Crops. Antalya, Turkey, May 22-26, 2006. Acta Horticulturae 781: 471–475. 170. LAUTERER P., MALENOVSKÝ I. & STRAKA J. 2008: Øasnokøídlí (Strepsiptera) Jizerských hor a Frýdlantska. (Strepsiptera of the Jizerské hory Mts. and Frýdlant region (northern Bohemia, Czech Republic)). Sborník Severoèeského muzea, Pøírodní vìdy 26: 89–93 (in Czech, English summary). 171. SELJAK G., MALENOVSKÝ I. & LAUTERER P. 2008: New records of jumping plant-lice from with description of Bactericera lyrata sp. n. (Hemiptera: Psylloidea). Revue Suisse de Zoologie 115: 527–540. 172. TEDESCHI R., LAUTERER P., BERTIGNONO L. & ALMA A. 2008: Hawthorn psyllid fauna in northwestern Italy. Bulletin of Insectology 61: 143–144. 173. ALMA A., BOCCA M., ÈERMÁK V., CHEN P.-P., D’URSO V., EXNEROVÁ A., GOULA M., GUGLIELIMINO A., KUNZ G., LAUTERER P., MALENOVSKÝ I., MAZZOGLIO P. J., NICOLI ALDINI R., OUVRARD D., †REMANE R., RINTALA T., SELJAK G., SÖDERMAN G., SOULIER-PERKINS A., ŠTYS P., TALVELLA L., TEDESCHI R. & WILSON M. 2009: Insecta Hemiptera collected in the Mont Avic Natural Park (Aosta Valley, Northwest Italy). Revue Valdôtaine d’Histoire Naturelle 63: 109–124. 174. ALMA A., CHEN P.-P., D’URSO V., GUGLIELMINO A., HOLLIER J., KUNZ G., LAUTERER P., MALENOVSKÝ I., MAZZOGLIO P. J., NICKEL H., ALDINI R. N., RINTTALA T., SELJAK G., SEYRING M., SÖDERMAN G., WILSON M. & WITSACK W. 2009: Auchenorrhyncha collected in the Canavese district (Northwest Italy) (Hemiptera, Auchenorrhyncha). Cicadina 10: 119–125. 175. BURCKHARDT D. & LAUTERER P. 2009a: Faunistic and taxonomic notes on jumping plant-lice from the Alps (Hemiptera: Psylloidea: ). Contributions to Natural History 12: 341–348. 176. BURCKHARDT D. & LAUTERER P. 2009b: Taxonomy of psyllids (Hemiptera, Psylloidea) associated with apple and stone fruits in Central and Southern . Mitteilungen der Schweizerischen Entomologischen Gesellschaft 82: 253–257. 177. FIALOVÁ R., NAVRÁTIL M., LAUTERER P., ÈERMÁK V. & Š AFÁØOVÁ D. 2009: Phytoplasma status of Cacopsylla pruni from apricot orchards and overwintering sites in Moravia. P. 46. In: ŠAFRÁNKOVÁ I. & ŠEFROVÁ H. (eds.): XVIII. èeská a slovenská konference o ochranì rostlin. 2.–4. záøí 2009. Sborník abstraktù. [XVIIIth Czech and Slovak Plant Protection Conference. 2–4th September 2009. Book of abstracts.] Mendelova zemìdìlská a lesnická univerzita, Brno, 239 pp. 178. JARAUSCH B., BURCKHARDT D., LAUTERER P. & J ARAUSCH W. 2009: Psyllids (Hemiptera: Psylloidea) captured in commercial apple and stone fruit orchards in southwest Germany, eastern France and northwest Switzerland. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 82: 205–215.

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179. LAUTERER P. 2009a: First record of Tomosvaryella rossica (Diptera: Pipunculidae) from the Czech Republic. Folia Faunistica Slovaca 14: 99–100. 180. LAUTERER P. 2009b: Pipunculidae Walker, 1834. JEDLIÈKA L., KÚDELA M. & STLOUKALOVÁ V. (eds.): Checklist of Diptera of Czech Republic and Slovakia. Electronic version 2. Available online: http://zoology.fns.uniba.sk/diptera2009 + CD-ROM (ISBN 978-80-969629-4-5). 181. MÜHLETHALER R., BURCKHARDT D., LAUTERER P. & NAGEL P. 2009: Taxonomy and biogeography of Central European Kybos (Insecta, Hemiptera, Cicadellidae). Deutsche Entomologische Zeitschrift 56: 15–40. 182. NAVRÁTIL M., VÁLOVÁ P., FIALOVÁ R., LAUTERER P., ŠAFÁØOVÁ D. & STARÝ M. 2009: The incidence of stolbur disease and associated yield losses in vegetable crops in South Moravia (Czech Republic). P. 58. In: ŠAFRÁNKOVÁ I. & ŠEFROVÁ H. (eds.): XVIII. èeská a slovenská konference o ochranì rostlin. 2. –4. záøí 2009. Sborník abstraktù. [XVIIIth Czech and Slovak Plant Protection Conference. 2–4th September 2009. Book of abstracts.] Mendelova zemìdìlská a lesnická univerzita, Brno, 239 pp. 183. TEDESCHI R., LAUTERER P., BRUSETTI L., TOTA F. & ALMA A. 2009: Composition, abundance and phytoplasma infection in the hawthorn psyllid fauna of northwestern Italy. European Journal of Plant Pathology 123: 301–310. 184. JARAUSCH W., JARAUSCH B., PECCERELLA T., DOLIT C. & LAUTERER P. 2010: Development of specific primers for the molecular identification of Cacopsylla picta, the main vector of apple proliferation disease. P. 57. In: BERTACCINI A., LAVIÑA A. & TORRES E. (eds.): Current status and perspectives of phytoplasma disease research and management. Abstract book of the combined meeting of Work Groups 1–4. February 1st and 2nd 2010, Sitges, . COST, European Cooperation of Science and Technology, 135 pp. Available online: http://costphytoplasma.eu/PDF%20files/WG%20BookwithiISBN.pdf 185. LAUTERER P. & DOROW W. H. O. 2010: Die Blattflöhe (Psylloidea) des Naturwaldreservats Goldbachs- und Ziebachsrück (Hessen), Untersuchungszeitsraum 1994–1996. In: DOROW W. H. O., BLICK T., KOEPELKE J. P. (eds.): Naturwaldreservats in Hessen. Band 11/2.2: Goldbachs- und Ziebachsrück. Zoologische Untersuchungen. 1994–1996. Teil 2. Mitteilungen der Hessischen Landesforstverwaltung 46: 99–110. 186. MALENOVSKÝ I. & LAUTERER P. 2010: Additions to the fauna of planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) in the Czech Republic. Acta Musei Moraviae, Scientiae biologicae 95(1): 49–122. 187. TEDESCHI R., LAUTERER P., BRUSETTU L., TOTA F., NARDU F. & ALMA A. 2010: Role of wild plants in epidemiology of fruit tree phytoplasmas and in ecology of the insect vectors. The case of hawthorn plants. P. 69. In: BERTACCINI A., LAVIÑA A. & TORRES E. (eds.): Current status and perspectives of phytoplasma disease research and management. Abstract book of the combined meeting of Work Groups 1–4. February 1st and 2nd 2010, Sitges, Spain. COST, European Cooperation of Science and Technology, 135 pp. Available online: http://costphytoplasma.eu/PDF%20files/WG%20BookwithiISBN.pdf 188. GUGLIELMINO A., LAUTERER P. & BÜCKLE C. 2011: Eupteryx cytinsularis, a new species of the melissae group (Rhynchota Auchenorrhyncha Cicadellidae) from Sicily, Sardinia and Corsica. Bulletin of Insectology 64: 23–26. 189. HOLÝ K., MACEK J. & LAUTERER P. 2011: Occurrence of Ibaliidae (Hymenoptera: Cynipoidea) in the Czech Republic. Sborník Severoèeského muzea, Pøírodní vìdy 29: 201–210. 190. LAUTERER P. 2011a: Psyllids (Hemiptera, Psylloidea) inhabiting the canopy and shrub layer of an oak and hornbeam forest in Báb near Nitra (Slovakia). Folia Faunistica Slovaca 16: 77–76. 191. LAUTERER P. 2011b: First finding of Porphyrophora polonica (Hemiptera: Coccoidea: ) in Slovakia. Folia Faunistica Slovaca 16: 77–78. 192. LAUTERER P., BARTOŠ R. & MILONAS P. 2011: First records of the jumping plant-louse jamatonica (Kuwayama) (Hemiptera: Sternorrhyncha: ) in Slovakia and Greece. Plant Protection Science 47: 40–43. 193. LAUTERER P. & LUDVÍKOVÁ H. 2011: Škùdce ovocných stromù ostnohøbetka ovocná již v Èechách. [Buffalo tree-hopper, a pest of fruit recorded in Bohemia.] Zahradnictví 2011(4): 12–14 (in Czech). 194. LAUTERER P., MALENOVSKÝ I. & ŠPRYÒAR P. 2011a: Šíøení invazní ostnohøbetky Stictocephala bisonia (Hemiptera: Cicadomorpha: Membracidae) v Èeské republice. P. 129. In.: BRYJA J., ØEHÁK Z. & ZUKAL J. (eds.): Zoologické dny Brno 2011. Sborník abstraktù z konference 17.–18. února 2011. Ústav biologie obratlovcù AV ÈR, Brno, 282 pp.

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195. LAUTERER P., MALENOVSKÝ I. & ŠPRYÒAR P. 2011b: Invasion of the Stictocephala bisonia (Hemiptera: Cicadomorpha: Membracidae) into the Czech Republic: current distribution and first records from Bohemia. Klapalekiana 47: 67–73. 196. LUDVÍKOVÁ H., LAUTERER P., SUCHÁ J. & FRÁNOVÁ J. 2011: Monitoring of psyllid species (Hemiptera, Psylloidea) in apple and pear orchards in East Bohemia. Bulletin of Insectology 64(Suppl.): 121–122. 197. LUDVÍKOVÁ H., SUCHÁ J. & LAUTERER P. 2011: Sledování výskytu mer rodu Cacopsylla ve výsadbách jabloní a hrušní ve východních Èechách. Zahradnictví 2011(10): 12–14. 198. ŠAFÁØOVÁ D., LAUTERER P., KORBASOVÁ Z., STARÝ M., VÁLOVÁ P. & NAVRÁTIL M. 2011: Hyalestehes obsoletus, a vector of stolbur phytoplasma: current situation in South Moravia, Czech Republic. Bulletin of Insectology 64(Suppl.): 127–128. 199. FRANOVA J., JAKESOVA H., KUBELKOVA D., KOLAROVA K., ORSAGOVA M., PETRZIK K. & LAUTERER P. 2011: Research of and phytoplasmas infecting clover plants in the Czech Republic. P. 51. In: Navas- Castillo J. & Fiallo-Olivé E. (eds.): 4th Conference of the International Working group on Legume and Vegetable Viruses (IWGLVV). Book of abstracts. Anteqera, Málaga, Spain, May 17–20, 2011. Instituto de Horticultura, subtropical y mediterránea „La Mayora“ (IHSM), Instituto Nacional de Investigatión y Tecnologia Agraria y Alimentaria, 121 pp. Available online: http://www.eelm.csic.es/iwglvv/files/ 4thIWGLVV-Book_of_abstracts.pdf 200. MALENOVSKÝ I. & LAUTERER P. 2012a: Jumping plant-lice (Hemiptera: Psylloidea) of the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). In: MALENOVSKÝ I., KMENT P. & KONVIÈKA O. (eds.): Species inventories of selected insect groups in the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). Acta Musei Moraviae, Scientiae biologicae (Brno) 96(2): 105–154. 201. MALENOVSKÝ I. & LAUTERER P. 2012b: Leafhoppers and planthoppers (Hemiptera: Auchenorrhyncha) of the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). In: MALENOVSKÝ I., KMENT P. & KONVIÈKA O. (eds.): Species inventories of selected insect groups in the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). Acta Musei Moraviae, Scientiae biologicae (Brno) 96(2): 155–322. 202. MALENOVSKÝ I., LAUTERER P., LABINA E. & BURCKHARDT D. 2012: Jumping plant-lice (Homoptera: Psylloidea) of Afghanistan. Acta Entomologica Musei Nationalis Pragae 52: 1–22. 203. LAUTERER P. 2013a: Køísi. [Leafhoppers and planthoppers.] Pp. 307–308. In: KARPAŠ R., VIŠÒÁK R. & VONIÈKA P. (eds.): Jizerské hory. O rašeliništích, kvìtenì a zvíøenì. [Jizerské hory Mts. On peatbogs, flora and fauna.] Nakladatelství RK, Liberec, 440 pp (in Czech). 204. LAUTERER P. 2013b: Mery. [Jumping plant-lice.] Pp. 308. In: KARPAŠ R., VIŠÒÁK R. & VONIÈKA P. (eds.): Jizerské hory. O rašeliništích, kvìtenì a zvíøenì. [Jizerské hory Mts. On peatbogs, flora and fauna.] Nakladatelství RK, Liberec, 440 pp (in Czech).

Igor Malenovský

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 19

ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 21–33, 2013

The oldest jumping plant-louse (Hemiptera: Sternorrhyncha) with comments on the classification and nomenclature of the Palaeogene Psylloidea

DAVID OUVRARD 1, DANIEL BURCKHARDT 2 & DALE GREENWALT 3 1 Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, U.K.; e-mail: [email protected] 2 Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland; e-mail: [email protected] 3 Paleobiology Department, National Museum of Natural History, Washington, DC, 20013, USA; e-mail: [email protected]

OUVRARD D., BURCKHARDT D. & GREENWALT D. 2013: The oldest jumping plant-louse (Hemiptera: Sternorrhyncha) with comments on the classification and nomenclature of the Palaeogene Psylloidea. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 21–33. – An Eocene compression fossil from the oil shales of the Kishenehn Formation in north-western Montana, USA, is described as Eogyropsylla paveloctogenarius sp.nov. It is the oldest known representative of the superfamily Psylloidea sensu stricto and the first Eocene (Lutetian, ca. 46 million year old) psyllid (: Aphalarinae) from the New World. The world psyllid Paleogene fauna is reviewed, and the following nomenclatural acts and changes in the classification are proposed: Carsidarina Bekker-Migdisova, 1985 = Palaeoaphalara Klimaszewski, 1993 syn.nov.; Proeurotica Bekker-Migdisova, 1985 = Plesioaphalara Klimaszewski, 1993 syn.nov.; Catopsylla Scudder, 1890 = Psyllites Cockerell, 1915 syn.nov.; Paleopsylloidini Bekker-Migdisova, 1985 = Palaeoaphalarinae Klimaszewski, 1993 syn.nov.; Parascenia Klimaszewski, 1997 stat. rev., removed from synonymy with Colposcenia Enderlein, 1929; Carsidarina Bekker-Migdisova, 1985 transferred from the to the Paleopsylloidini Bekker-Migdisova, 1985 (Aphalarinae); Catopsylla Scudder, 1890 transferred from the Psyllidae to the Paleopsylloidini. Keywords. Aphalarinae, Eogyropsylla, jumping plant-lice, psyllids, fossil, biogeography, palaeontology, systematics, taxonomy, Holarctic Region, Kishenehn Formation

Introduction Fossils provide a direct way of character polarisation when inferring phylogenies (OUVRARD et al. 2010). Fossils also document the diversification of morpho-anatomical structures through time, and help dating the age of lineages (HARBACH & GREENWALT 2012). A major problem with fossils is, however, that they are often incompletely preserved and that taxonomically important characters cannot be observed. This is also true in the jumping plant-lice or psyllids (Hemiptera: Sternorrhyncha: Psylloidea) where phylogenetically relevant characters are often difficult to observe (e.g., larval tarsal arolium) and homoplasy is important (e.g., presence/absence of a pterostigma in the forewing) (BURCKHARDT & QUEIROZ 2013).

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Psyllids are small sap-sucking insects distributed worldwide. Most of the over 3850 described species (LI 2011) develop exclusively on dicotyledonous plants. OUVRARD et al. (2010) showed that Psylloidea sensu stricto are mainly characterised by the autapomorphic highly specialised metacoxa, with a striking development of its meron, on which the jumping muscles attach. The closest known lineages, without specialised coxa, forming the stem-group of extant Psylloidea are the extinct families Malmopsyllidae and Liadopsyllidae. The youngest known representative of Liadopsyllidae, Liadopsylla hesperia Ouvrard et Burckhardt, 2010, was described from the Upper (Turonian) and is ca. 90 million years (myr.) old. After the extinction of the stem-group in the Late Cretaceous, the fossil record of Psylloidea sensu lato shows a gap of more than 46 myr. Fossil Psylloidea with modified coxae appear first in Baltic amber (Eocene). This gap covers almost exactly the time of the angiosperm radiation during the Upper Cretaceous and Paleocene. The simultaneous rarefaction in the psyllid fossil record, the explosion of flowering plant diversity and the appearence of modified morphological and biological traits in modern psyllids show how drastically the evolution and diversification of new plant families has influenced the evolution of Psylloidea. Together with the modification of the meral part of the metacoxa fused with the complex metathoracic furcae, the observed modified characters are the shortened rostrum, the robust femora, and the distally enlarged metatibiae bearing apical sclerotised spurs (OUVRARD et al. 2010). The oldest described Psylloidea sensu stricto were found in Baltic amber dating from the Middle (44 myr.) to Upper Eocene (38 myr.) (DROHOJOWSKA & SZWEDO 2011; Tables 1 and 2). These are the genera Eogyropsylla Klimaszewski, 1993, Paleopsylloides Bekker-Migdisova, 1985 and Parascenia Klimaszewski, 1997 (= Colposcenia Enderlein, 1929) which were assigned to the Paleopsylloidini (Aphalaridae: Aphalarinae) (KLIMASZEWSKI 2001), as well as Protoscena Klimaszewski, 1997, assigned to the Paurocephalinae (Rhinocolidae) by KLIMASZEWSKI (1997b). KLIMASZEWSKI (2001) defined the tribe Paleopsylloidini as follows: head lacking genal processes; vertex longer than half as wide; antenna 10-segmented, about as long as forewing width, with a subapical rhinarium on each of segments 4–9; clypeus subglobular; pronotum subrectangular; forewing with membranous pterostigma; hindwing with R+M; and male proctiger one-segmented with wing-like posterior processes. Considerably younger, from late Eocene or early Oligocene, are the genera Carsidarina Bekker-Migdisova, 1985, Proeurotica Bekker-Migdisova, 1985, Lapidopsylla Klimaszewski, 1993, Plesioaphalara Klimaszewski, 1993 and Palaeoaphalara Klimaszewski, 1993 which are all known from compression fossils from the Isle of Wight, UK. The first genus was referred to Carsidaridae (BEKKER-MIGDISOVA 1985), the last to the Palaeoaphalarinae (Aphalaridae), and the rest to the Paleopsylloidini (KLIMASZEWSKI 2001). The Palaeoaphalarinae were separated from Aphalarinae by the two-segmented male proctiger. The youngest Paleogene genera, at the Eocene/Oligocene border, are the three genera Catopsylla Scudder, 1890, Necropsylla Scudder, 1890 and Psyllites Cockerell,

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1915 from Florissant, Colorado, USA, referred to the Psyllidae, Paleopsylloidini and , respectively (BEKKER-MIGDISOVA 1985, EVANOFF et al. 2001). The Paleogene Psylloidea (Tables 1 and 2) have been referred to extinct genera of the extant families Aphalaridae, Calophyidae, Carsidaridae and Psyllidae (BEKKER- MIGDISOVA 1985, SHCHERBAKOV & POPOV 2002), unlike those from the Neogene which are morphologically close to or congeneric with extant genera (BURCKHARDT & OUVRARD 2012). In Dominican amber (Miocene) all extant families are represented except for Carsidaridae and (BURCKHARDT & OUVRARD 2012; D. Burckhardt, unpublished data). Here we describe an Eocene compression fossil from the oil shales of Kishenehn Formation in north-western Montana, USA which we refer to Eogyropsylla. The attribution to a particular genus or even subfamily or family is difficult as many taxonomically relevant characters are missing and the systematics of Paleogene psyllids is in urgent need of revision. The new species is the oldest known representative of the superfamily Psylloidea sensu stricto and the first Eocene psyllid from the New World. In addition, known Paleogene taxa are discussed and their classification is reviewed.

Materials and methods One female specimen was collected in 2012 at the Constenius Park site along the Middle Fork of the Flathead River under the auspices of United States Forest Service Permit HUN281. The specimen is deposited at the National Museum of Natural History in Washington, D.C. (USNM 553523). The oil shale of the Middle Sequence of the Coal Creek Member of the Kishenehn Formation has been dated to 46.2 ± 0.4 million years old by 40Ar/39Ar analysis and 43.5 ± 4.9 million years old by fission-track analysis (CONSTENIUS 1996). The age estimates are taken from the chronostratigraphic chart published by the International Commission on Stratigraphy in August 2012 (available at http://www.stratigraphy.org/). The observation and photography were made after immersion of the specimen in 96% ethanol. Pictures were taken using a Canon EOS 50 D camera with a Macrolens MP- E 65 mm f/2.8 1–5X. Images were viewed with the software Canon EOS utility and stacked images were combined using the software Combine Z5. The updated taxonomy together with literature and geological ages for fossils for all cited taxa can be found on the Psyl’list online database (OUVRARD 2013). Morphological terminology follows HOLLIS (2004).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 23 D. OUVRARD ET AL.

Table 1. Paleogene Psylloidea according to the classifications of BEKKER-MIGDISOVA (1985) and KLIMASZEWSKI (1997b, 2001).

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Taxonomy Eogyropsylla paveloctogenarius sp.nov. (Fig. 1) Type locality. Park Middle Fork of the Flathead River 2 miles south of Stanton Creek, Montana, USA (W 113° 42.085′, N 48°23.517′). Type material. Holotype: female, National Museum of Natural History in Washington, D. C. (USNM no. 553523). Type stratum. Middle Eocene, Middle Sequence of the Coal Creek Member, Kishenehn Formation (Lutetian). Description. Adult, female. Body dark brown. Legs and forewing veins lighter brown. Thorax robust, strongly arched. Forewing ovoid, pterostigma absent, veins R and

M+Cu subequal, vein R1 long, almost perpendicular to vein Rs, cell cu1 long and narrow. Abdomen large and hardly constricted at base. Metatibia relatively strong, barely expanding apically. Metabasitarsus enlarged. Apical tarsomere of hind leg bearing strong claws. Head. In very bad condition, so that no constitutive part is visible with exception of the antennae and a portion of one eye. Nothing can be said about the shape of the genae nor the vertex. Antennae almost as long as forewing width, with two strongly enlarged basal articles. Segmentation of the flagellum not visible. Apical part slightly enlarged. Thorax with sclerites visible from the side (laterally) caudally, but more from above frontally, because of a probable twisting of the anterior part of the body. Pronotum not visible. Mesopraescutum and mesoscutum large, subequal in length. Ventrally, part of the mesopleuron is visible, but strongly damaged. Second peritreme visible, as well as part of the pleural suture in mesothorax. Metathorax not visible, not distinguishable from the anterior segments of the abdomen. Legs. Coxae not visible. Only apical portion of one hind leg visible. Tibia relatively long, more or less evenly thick throughout entire length, slightly thicker distally. Tarsus two-segmented, basal segment thick, expanded apically, distal segment ovoid, probably bearing two strong claws (one visible). Forewing oblong-oval; 1.9 mm long, 0.9 mm wide. Only left forewing flat, as shown by the clearly visible and unfolded large anal cell (right wing probably slightly deformed

Table 2. Paleogene Psylloidea arranged according to geological epoch and stage.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 25 D. OUVRARD ET AL.

Fig. 1. Female holotype of Eogyropsylla paveloctogenarius sp.nov. (USNM no. 553523).

during fossilization). Widest in apical third, without pterostigma nor visible thickening of vein C. Costal break present. Cell r1 broad. Vein R1 almost perpendicular to anterior wing edge, forming a right angle with vein Rs. Vein Rs sinuous and upturned at apex. Veins R and M+Cu subequal in length. Vein M hardly arched, vein M1+2 almost straight, in the prolongation of M, ending approximately at wing apex. Vein Cu 0.3 mm long. Cell cu1 elongate, 2.45 times as long as high. Anal break present, adjacent to merging point of vein

Cu1b with posterior margin of the wing. A nodal line seems to be developed. Hindwing not preserved. Abdomen ovoid. Three large dark-brown distal tergites visible. Female terminalia cuneate. Proctiger 0.4 mm long, with straight dorsal margin. Anus 0.14 mm long. Male. Unknown.

26 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Oldest jumping plant-louse with comments on Palaeogene Psylloidea

Measurements. In µm. Total body length 1747, antenna length 630 (left) – 705 (right), forewing length 1906 (left) – 1942 (right), forewing width 936 (left) – 822 (right), vein

R length 424, vein M+Cu length 406, vein M length 604, vein M1+2 length 486 (left) –

605 (right), vein M3+4 length 443 (left) – 441 (right), vein Cu length 294, cell Cu1 length

617 (left) – 654 (right), cell Cu1 height 252 (left) – 189 (right), abdomen length 1087, proctiger length 419, anus diameter 137. Distribution. USA (Montana). Etymology. Named in honour of our friend and colleague Dr. Pavel Lauterer on the occasion of his 80th birthday. Noun in apposition. Differential diagnosis. Eogyropsylla paveloctogenarius sp.nov. is closest to E. parva

Klimaszewski, 1997. The latter has a smaller body size with a longer cell m1 and veins

Rs and R1 forming an acute angle in forewing, and with elongated female terminalia. The former differs from all other Eogyropsylla species in the absence of a pterostigma, the more rounded forewing shape, the elongated cell cu1 and the cuneate female terminalia.

Discussion Compared to other insects, the fossil record of psyllids is scarce in general and completely lacking for the period from the late Upper Cretaceous (Coniacian) to the early Eocene (Ypresian). This paucity could be due to the minute body size of jumping plant- lice which can easily be overlooked during stone examinations but, more likely, it reflects a genuine rarity of psyllids during the Upper Cretaceous and the Palaeocene due to the profound changes in floras and climates bringing about the extinction of Liadopsyllidae and Malmopsyllidae followed by the development of a new group, the recent Psylloidea (KLIMASZEWSKI 1997a, 2001). The discovery of Eogyropsylla paveloctogenarius sp.nov. from Lutetian deposits is thus important, as it is the oldest representative of extant Psylloidea and it is the first New World psyllid known from the Eocene. Until now only two North American imprints of Aphalarinae were known from the Oligocene: Necropsylla rigida Scudder, 1890, and Necropsylla rigidula Cockerell, 1911, both from Florissant, Colorado, USA (the latter described from a very small forewing fragment). Two additional species from other families were described from the same stratum: Psyllites crawfordi Cockerell, 1915 (Calophyidae) and Catopsylla prima Scudder, 1890 (Psyllidae). Eogyropsylla paveloctogenarius is placed in Eogyropsylla as it resembles the other species in the general body shape, the forewing venation and antennal length; other characters cannot be judged as they are not clearly visible. It differs from the remaining species in the lack of a pterostigma. Even though the presence or absence of a pterostigma is often stable within a genus, there are exceptions. In Gyropsylla (Aphalaridae, Aphalarinae) two species bear a membranous pterostigma, G. cannela (Crawford, 1925) and G. ilicis (Ashmead, 1881), and four lack one: G. chiriquiensis Brown et Hodkinson, 1988, G. clypeata Brown et Hodkinson, 1988, G. longilabiata (Li, 1991) and G. speggazziniana (Lizer, 1919). Within Eogyropsylla, E. paveloctogenarius

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 27 D. OUVRARD ET AL. resembles most E. parva in the wing shape and dimensions. It differs from E. parva in the absence of a pterostigma and shorter female terminalia. KLIMASZEWSKI (1993) and DROHOJOWSKA (2011) recorded a well-developed pterostigma in Eogyropsylla species. The forewing of E. parva illustrated by KLIMASZEWSKI (1997a) lacks, however, a pterostigma but this is an error (holotype in Naturhistorisches Museum Basel, examined). KLIMASZEWSKI (1993) placed Eogyropsylla in the extinct tribe Paleopsylloidini and suggested a close relationship to the extant genus Gyropsylla Brèthes, 1921. Eogyropsylla differs from Paleopsylloides in the forewing veins R and M+Cu which are of subequal length, rather than vein M+Cu distinctly longer that vein R. The latter state is relatively rare in psyllids and occurs for instance in many . Later, KLIMASZEWSKI (1997a) placed Eogyropsylla in the tribe Gyropsyllini. As stated by DROHOJOWSKA (2011), Eogyropsylla does, however, not share any derived character with Gyropsylla, and its phylogenetic relationship remains doubtful. Based on the presence of posterior processes on the male proctiger in Paleopsylloides and Parascenia their assignment to the subfamily Aphalarinae is well-supported. Eogyropsylla, only known from females and one last instar larva, resembles strongly the latter genus suggesting a close relationship. The monophyly of the tribe Paleopsylloidini is questionable as it is defined by likely symplesiomorphies and homoplasies (DROHOJOWSKA 2011). The tribe may be paraphyletic with respect to extant Aphalarinae whose internal phylogenetic relationships are also in need of revision (BURCKHARDT & OUVRARD 2012). DROHOJOWSKA (2011) suggested also that characters defining Eogyropsylla are symplesiomorphies, and Parascenia and Paleopsylloides are monotypic. The last differs from the first two genera in the wing venation but the first two are similar and may represent females and males of the same genus. KLIMASZEWSKI (1997a) stated that Parascenia is similar to Colposcenia in the enlarged lobes of the male proctiger, and BURCKHARDT & OUVRARD (2012) synonymised Parascenia with Colposcenia. Enlarged lobes occur also in other extant aphalarine genera such as Caillardia, and Parascenia differs from Colposcenia in the less inclined head which is angular rather that rounded anteriorly. More work is needed to confirm that Eogyropsylla and Parascenia are synonyms. We therefore suggest to reinstate the latter as Parascenia stat. rev. in the Paleopsylloidini. According to KLIMASZEWSKI (1997b) Protoscena lacks meracanthi. For this reason BURCKHARDT & OUVRARD (2012) assigned the genus to the Rhinocolinae (Aphalaridae). The placement of the other Paleogene genera, known from impressions only, is difficult to judge as they consist often of single forewings. From the Isle of Wight, UK, compression fossils have been attributed to the genera Carsidarina, Proeurotica, Lapidopsylla, Palaeoaphalara and Plesioaphalara. Carsidarina and Palaeoaphalara share the short pterostigma, the short vein Cu1 and the large cells m1 and cu1. We consider these characters as synapomorphies and suggest that the two are synonyms. The former was referred to the Carsidaridae (BEKKER-MIGDISOVA 1985) whose forewings differ considerably in the usually acute or subacute apex (except Epicarsa) (HOLLIS 1987) and the very small cell cu1. Epicarsa is characterised by Rs and M1+2 which are in broad contact. Palaeoaphalara was referred to the subfamily Palaeoaphalarinae (Aphalaridae)

28 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Oldest jumping plant-louse with comments on Palaeogene Psylloidea on the basis of the two-segmented proctiger (KLIMASZEWSKI & POPOV 1993). Providing this observation is correct, it is probably a primitive character and unsuitable for phylogenetic reconstruction. The Palaeoaphalarinae was treated as incertae sedis by BURCKHARDT & OUVRARD (2012). It appears that Paleopsylloidini and Palaeoaphalarinae refer to the same group and we synonymise them and transfer Carsidarina to the Paleopsylloidini. Again there is a striking resemblance between the forewings of

Proeurotica and Plesioaphalara in the small cell m1 and very long cell cu1, suggesting that they are congeneric. We propose here their synonymy. Among the youngest Paleogene genera Catopsylla, Necropsylla and Psyllites from Florissant, Colorado, USA, the first and last share an apically broadly rounded forewing with large m1 and cu1 cells indicating close relationship and synonymy of the two genera. They were attributed to the Psyllidae and Calophyidae, respectively, for neither of which is there any evidence. As the forewing characters resemble those of Carsidarina the taxon can be accommodated in the Paleopsylloidini. Among Paleogene psyllids, Eogyropsylla and Necropsylla have several characters in common suggesting that they are more closely related to each other than either of them is to Paleopsylloides. Their type-species, viz. E. eocenica Klimaszewski, 1993 and N. rigida, share the following features: forewing oblong-oval, Rs long, cell r1 very large, R and M+Cu of subequal length, M and M1+2 forming a relatively straight line, cell cu1 elongate, and female terminalia cuneate. Eogyropsylla and Necropsylla differ from each other in the presence of a pterostigma in the former and a vein R2 in the latter where the angle between R1 and Rs is more acute. Unlike the Nearctic taxa, the Palaearctic Eogyropsylla species are all from Baltic amber (DROHOJOWSKA 2011) where much more morphological detail is visible than in compression fossils. There is also some doubt whether the pterostigma of Eogyropsylla and the presence of the vein R2 in Necropsylla are in fact the same and just an artefact due to the differences in fossilisation (amber versus compression fossil). The assignment of E. paveloctagenarius to Eogyropsylla is supported by the similar age of these taxa, all from the Eocene. As reported by DROHOJOWSKA & SZWEDO (2011) the precise dating of Baltic amber is difficult and varies from Middle Eocene (ca. 44 myr. old) to Upper Eocene (ca. 38 myr. old). The Nearctic Necropsylla is considerably younger with an age difference of about 7 myr. Regarding the biogeography, the newly described Eogyropsylla species extends considerably the distribution area of the genus, which was only known from the Old World until now. According to HODKINSON (1980), the development of a common fauna between Nearctic and the western Palaearctic, as pointed out also by KLIMASZEWSKI (1987), occurred until the final separation of the two regions, when the Atlantic corridor definitely closed in the Middle Eocene (ca. 40 myr. ago), which corresponds to the approximate age of our fossil and its congeneric species from Baltic amber. It means that the members of the genus in North America and the members in Europe had not been separated for a long time before their extinctions, which is corroborated by the fact that they do not show striking divergence in wing morphology, except for the absence of a pterostigma. Among the extant aphalarine genera, Aphalara Foerster, 1848 and Craspedolepta Enderlein, 1921, lack a pterostigma, and both have a Holarctic

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 29 D. OUVRARD ET AL. distribution. Other Palaearctic genera containing species without a pterostigma are Brachystetha Loginova, 1964, Crastina Loginova, 1964, Rhombaphalara Loginova, 1964, and Xenaphalara Loginova, 1961. The following monotypic aphalarine genera restricted to the New World all have a pterostigma: Limataphalara Hodkinson, 1992; Neaphalara Brown et Hodkinson, 1988; Hodkinsonia Burckhardt, Espírito-Santo, Fernandes et Malenovský, 2004; and Lanthanaphalara Tuthill, 1959. The New World/Asian genus Gyropsylla which is associated with Ilex spp. (Aquifoliaceae) contains two described species with and four lacking a pterostigma. Being considered as an apomorphic state by KLIMASZEWSKI (2001), the lack of a pterostigma occurred probably several times independently all along the course of evolution of the Aphalarinae, and is hence of little phylogenetic significance. According to HODKINSON (1980, 1992) the Neotropical aphalarine genera, apart from Gyropsylla, all monotypic, are relicts from a larger fauna, and the presence of Gyropsylla and Limataphalara in southern North America is due to more recent (less than 20 myr. old) northern migration from the Neotropics, together with other psyllid taxa. This biogeographic evidence is corroborated by the discovery of E. paveloctogenarius which does not share derived characters with Gyropsylla, contrary to the assumptions of KLIMASZEWSKI (1993, 2001) on which the name Eogyropsylla itself is based. This weakly supported relationship had already been impeded by the mention by KLIMASZEWSKI (1997b) of the affinity between Gyropsylla and Protoscena baltica (not a member of the Aphalarinae, but of the Rhinocolinae), which added confusion to the situation. As shown earlier, the lack of strong evidence for a close relationship between Eogyropsylla and Gyropsylla has already been mentioned by DROHOJOWSKA (2011). Eogyropsylla is probably part of the stem-group which gave rise to the extant Aphalarinae genera. However, contrary to the Palaearctic, no diversification occurred in the Nearctic region, where the lineage of E. paveloctogenarius became extinct. In Eurasia, on the contrary, the diversification of the other members of the genus has gone on, and a centre of diversification occurred around Central Asia. Some of the genera reached North Africa in a South-Western expansion (e.g., Caillardia De Bergevin, 1931, Colposcenia, Crastina, Rhodochlanis Loginova, 1964), Colposcenia and Crastina even in the Afrotropical Region, while some others spread out on an Eastern track to the Far East (Epheloscyta Loginova, 1976) or were even able to cross the Beringia region at some point (Aphalara and Craspedolepta) as proposed by HODKINSON (1978, 1980). The depositional environment of E. paveloctogenarius was both lacustrine and paludal; the exposures have an uncommonly large numbers of mosquito fossils (35 to date), two of which have recently been described (HARBACH & GREENWALT 2012). Other fauna described from the Formation include fishes, mollusks and vertebrates (CONSTENIUS et al. 1989, LI & WILSON 1994). The molluscan fauna of the Kishenehn Formation has been divided into four different successional groups, the first exemplified by Gastrocopta miniscula Pierce, 1992. Gastrocopta pellucida (Pfeiffer, 1840) selected as the extant analog of G. miniscula, currently lives in an environment characterized by a mean annual temperature of 25–27°C (PIERCE & CONSTENIUS 2001). The early arboreal primate Tarkadectes montanensis McKenna, 1990 originally described from the Coal Creek member (MCKENNA 1990), was recently assigned to the extinct family

30 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Oldest jumping plant-louse with comments on Palaeogene Psylloidea

Omomyidae (NI et al. 2010). Most closely related to the Omomyidae are the insectivorous Tarsiidae, extant species of which are restricted to islands of Southeast Asia (GROVES & SHEKELLE 2010). The temperatures of this period were significantly warmer than the present day and were likely wet subtropical in nature. Although fossil leaves are moderately common in layers of siltstone and sandstone from the Kishenehn Formation, only very preliminary reports have been published (ROSS 1959, CONSTENIUS et al. 1989). However, more detailed analyses of palynomorphs identified 27 different families of plants (HOPKINS & SWEET 1976), but none of the common host-plant families for extant Aphalarinae (mainly Amaranthaceae, Asteraceae, Onagraceae, Polygonaceae and Tamaricaceae) has been reported from the site. The presence of fossil pollen from cattail (Typha) and waterfern (Azolla) support deposition in a paludal environment while the presence of abundant pollen of the family Pinaceae indicate the existence of a surrounding forested highland.

Nomenclatorial acts Following new nomenclatorial acts are proposed here: Carsidarina Bekker-Migdisova, 1985 = Palaeoaphalara Klimaszewski, 1993 syn.nov. Proeurotica Bekker-Migdisova, 1985 = Plesioaphalara Klimaszewski, 1993 syn.nov. Catopsylla Scudder, 1890 = Psyllites Cockerell, 1915 syn.nov. Paleopsylloidini Bekker-Migdisova, 1985 = Palaeoaphalarinae Klimaszewski, 1993 syn.nov. Parascenia Klimaszewski, 1997 stat. rev., removed from synonymy with Colposcenia Enderlein, 1929 Carsidarina Bekker-Migdisova, 1985 transferred from the Carsidaridae to the Paleopsylloidini Bekker- Migdisova, 1985 Catopsylla Scudder, 1890 transferred from the Psyllidae to the Paleopsylloidini

Conclusion The newly described fossil jumping plant-louse supports GRIMALDI et al.’s (2002) observation that fossil insects from the Eocene belong almost exclusively to modern families, but not necessarily to modern genera. The exact relationships between the known Eocene fossils in Aphalarinae and the extant fauna are difficult to evaluate. The monophyly of Aphalarinae is reasonably well supported (DOBREANU & MANOLACHE 1960, BURCKHARDT 1987) but not that of its five constituent tribes (BURCKHARDT & OUVRARD 2012, BURCKHARDT & QUEIROZ 2013). In addition there are no derived characters which would group the extinct members of the subfamily together with either the New World Aphalarinae or the Palaearctic ones. Furthermore, important diagnostic characters for defining aphalarine genera are not visible on rock imprints, like characters relating to the vertex, clypeus, hindlegs or the male terminalia. However, in extending the distribution of the genus to North America, the discovery of E. paveloctogenarius sp.nov., which is also the oldest member of the entire superfamily, allows specification of more detailed evolutionary scenarios for Aphalarinae that still have to be tested in a phylogenetic framework.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 31 D. OUVRARD ET AL.

Acknowledgements The authors are grateful to Mr. Laurent Fauvre for his help in photographing the specimen and to Ms. Madeleine Paillé for fruitful discussions on the latin name. This is contribution number 282 of the Evolution of Terrestrial Ecosystems consortium at the National Museum of Natural History in Washington, D.C.

References

BEKKER-MIGDISOVA E. E. 1985: Iskopaemye nasekomye Psillomorfy. [Fossil Psyllomorpha]. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 206: 1–94 (in Russian). BURCKHARDT D. 1987: Jumping plant lice (Homoptera: Psylloidea) of the temperate Neotropical region. Part 1: Psyllidae (subfamilies Aphalarinae, Rhinocolinae and Aphalaroidinae). Zoological Journal of the Linnean Society 89: 299–392. BURCKHARDT D. & OUVRARD D. 2012: A revised classification of the jumping plant-lice (Hemiptera: Psylloidea). Zootaxa 3509: 1–34. BURCKHARDT D. & QUEIROZ D. L. 2013. Systematics of the Neotropical jumping plant-louse genus Limataphalara (Hemiptera: Psylloidea: Aphalaridae) and phylogenetic relationships within the subfamily Aphalarinae. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 35–56. CONSTENIUS K. N. 1996: Late Paleogene extensional collapse of the Cordilleran foreland fold and thrust belt. Geological Society of America Bulletin 108: 20–39. CONSTENIUS K. N., DAWSON M. R., PIERCE H. G., WALTER R. C. & WILSON M. V. H. 1989: Reconnaissance paleontologic study of the Kishenehn Formation, northwestern Montana and southeastern British Columbia. Pp. 189–203. In: FRENCH D. E. & GRABB R. F. (eds.): 1989 Field Conference Guidebook. Montana Centennial Edition. Geologic Resourses of Montana, Volume 1. Montana Geological Society, Billings, 422 pp. DOBREANU E. & MANOLACHE C. 1960: Recherches sur l’organe copulateur de quelques espèces de psylles (Psylloidea, Triozinae et Aphalarinae). Revue de Biologie 5: 87–99. DROHOJOWSKA J. 2011: Eogyropsylla sedzimiri sp. nov. from Eocene Baltic amber with a key to the species of the fossil genus Eogyropsylla Klimaszewski, 1993 (Hemiptera: Sternorrhyncha: Psylloidea). Zootaxa 2803: 41–48. DROHOJOWSKA J. & SZWEDO J. (2011) New Aleyrodidae (Hemiptera: Sternorrhyncha: Aleyrodomorpha) from Eocene Baltic amber. Polish Journal of Entomology 80: 659–677. EVANOFF E., MCINTOSH W. C. & MURPHEY P. C. 2001: Stratigraphic summary and 40Ar/39Ar geochronology of the Florissant Formation. Pp. 1–16. In: EVANOFF E., GREGORY-WODZICKI K. M. & JOHNSON K. R. (eds.): Fossil flora and stratigraphy of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature and Science Series 4, number 1: 1–69. GRIMALDI D. A., ENGEL M. S. & NASCIMBENE P. C. 2002: Fossiliferous Cretaceous amber from Myanmar (Burma): its rediscovery, biotic diversity, and paleontological significance. American Museum Novitates 3361: 35–37. GROVES C. & SHEKELLE M. 2010: The genera and species of Tarsiidae. International Journal of Primatology 31: 1071–1082. HARBACH R. E. & GREENWALT D. 2012: Two Eocene species of Culiseta (Diptera: Culicidae) from the Kishenehn Formation in Montana. Zootaxa 3530: 25–34. HODKINSON I. D. 1978: The psyllids (Homoptera: Psylloidea) of Alaska. Systematic Entomology 3: 333–360. HODKINSON I. D. 1980: Present day distribution patterns of the holarctic Psylloidea (Homoptera: Insecta) with particular reference to the origin of the nearctic fauna. Journal of Biogeography 7: 127–146. HODKINSON I. D. 1992: A new genus of psyllid (Homoptera: Psylloidea: Aphalaridae) from Florida. Florida Entomologist 75: 84–89. HOLLIS D. 1987: A review of the Malvales-feeding psyllid family Carsidaridae (Homoptera). Bulletin of the British Museum (Natural History), Entomology 56(2): 87–127. HOLLIS D. 2004: Australian Psylloidea: jumping plantlice and lerp insects. Canberra, Australia, 216 pp.

32 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Oldest jumping plant-louse with comments on Palaeogene Psylloidea

HOPKINS W. S. JR. & SWEET A. R. 1976: A microflora from a short section of the Paleogene Kishenehn Formation, Southeastern British Columbia. Geological Survey of Canada 76-1B: 307–309. KLIMASZEWSKI S. M. 1987: Relationships within the Aphalarinae Löw subfamily (Homoptera, Psylloidea). Polskie Pismo Entomologiczne 57: 211–233. KLIMASZEWSKI S. M. 1993: New species of Eocene psyllids representing the tribe Paleopsylloidini Becker- Migdisova (Homoptera, Psylloidea). Acta Biologica Silesiana 22: 9–18. KLIMASZEWSKI S. M. 1997a: New psyllids from the Baltic amber (Insecta: Homoptera, Aphalaridae). Mitteilungen aus dem Geologisch-Palaeontologischen Institut der Universität Hamburg 80: 157–171. KLIMASZEWSKI S. M. 1997b: Protoscena baltica gen. et sp. n. from the Eocene Baltic amber (Hemiptera, Homoptera). Annalen des Naturhistorischen Museums in Wien 98A: 69–72. KLIMASZEWSKI S. M. 2001: Jumping plant lice from the family Aphalaridae in Central Asian Province (Hemiptera: Sternorrhyncha: Psylloidea). Annals of the Upper Silesian Museum in Bytom, Entomology 10–11: 179–201. KLIMASZEWSKI S. M. & POPOV Y. 1993: New fossil hemipteran insects from Southern England (Hemiptera: Psyllina + ). Annals of the Upper Silesian Museum, Entomology Suppl. 1: 13–36. LI F. 2011: Psyllidomorpha of China (Insecta: Hemiptera). Science Press, Beijing, 1976 pp (in Chinese, English abstract). LI G.-Q. & WILSON M. V. H. 1994: An Eocene species of Hiodon from Montana, its phylogenetic relationships, and the evolution of the postcranial skeleton in the Hiodontidae (Teleostei). Journal of Vertebrate Paleontology 14: 153–167. MCKENNA M. C. 1990: Plagiomenids (Mammalia: ?Dermoptera) from the Oligocene of Oregon, Montana and South Dakota, and Middle Eocene of northwestern Wyoming. Pp. 211–234. In: BROWN T. M. & ROSE K. D. (eds.): Dawn of the age of mammals in the northern part of the Rocky Mountain interior, North America. Special Papers of the Geological Society of America 243: 1–241. NI X., MENG J., BEARD K. C., GEBO D. L., WANG Y. & LI C. 2010: A new tarkadectine primate from the Eocene of Inner Mongolia, China: phylogenetic and biogeographic implications. Proceedings of the Royal Society of London, Series B, Biological Sciences 277: 247–256. OUVRARD D. 2013: Psyl’list – The World Psylloidea Database. Available online from http://www.hemiptera- databases.com/psyllist (accessed on 11 June 2013). OUVRARD D., BURCKHARDT D., AZAR D. & GRIMALDI D. 2010: Non-jumping plant-lice in Cretaceous amber (Hemiptera: Sternorrhyncha: Psylloidea). Systematic Entomology 35: 172–180. PIERCE H. G. & CONSTENIUS K. N. 2001: Late Eocene – Oligocene nonmarine mollusks of the northern Kishenehn Basin, Montana and British Columbia. Annals of the Carnegie Museum 70: 1–112. ROSS C. P. 1959: Geology of Glacier National Park and the Flathead Region, Northwestern Montana. USGS Professional Paper, Report 296: 1–125. SHCHERBAKOV D. E. & POPOV Yu. A. 2002: Superorder Cimicidea Laicharting, 1781. Order Hemiptera Linné, 1758. The Bugs, , Plantlice, Scale Insects, etc. Pp. 143–157. In: RASNITSYN A. P. & QUICKE D. L. J. (eds.): History of Insects. Kluwer Academic, Dordrecht, 517 pp.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 35–56, 2013

Phylogenetic relationships within the subfamily Aphalarinae including a revision of Limataphalara (Hemiptera: Psylloidea: Aphalaridae)

DANIEL BURCKHARDT1 & DALVA LUIZ DE QUEIROZ2 1Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland; e-mail: [email protected] 2Embrapa Florestas, Estrada da Ribeira, km 111, C. postal 319, 83411-000, Colombo, PR, Brazil; e-mail: [email protected]

BURCKHARDT D. & QUEIROZ D. L. 2013: Phylogenetic relationships within the subfamily Aphalarinae including a revision of Limataphalara (Hemiptera: Psylloidea: Aphalaridae). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 35–56. – The hitherto monotypic Neotropical genus Limataphalara Hodkinson, 1992 is revised and three new species are described and illustrated: Limataphalara eucosma sp.nov. from Brazil associated with Nectandra lanceolata (Lauraceae), L. hollisi sp.nov. from Costa Rica associated with N. membranacea, and L. lautereri sp.nov. from Brazil associated with N. cuspidata. Based on the new species the generic diagnosis is revised and the presumed host association with Nectandra is confirmed. For the first time larvae of the genus are described and the phylogenetic relationships of all known genera within Aphalarinae are analysed with cladistic methodology using 16 morphological characters. The analysis resulted in three most parsimonious trees which differ considerably from previously published phylogenetic schemes resulting in the following new family level synonymies: Aphalarinae Löw, 1879 (= Caillardiini Loginova, 1964, syn.nov.; Coelocarinae Li, 2011, syn.nov.; Colposceniini Bekker-Migdisova, 1973, syn.nov.; Craspedoleptini Klimaszewski, 2001, syn.nov.; Eumetoecini Li, 2011, syn.nov.; Gyropsyllini White & Hodkinson, 1985, syn.nov.; Rhombaphalarini Klimaszewski, 1987, syn.nov.; Stigmaphalarini Vondráèek, 1957, nomen nudum, syn.nov.; Xenaphalarini Loginova, 1964, syn.nov.), and the following new genus level synonymy: Rhodochlanis Loginova, 1964 (= Rhombaphalara Loginova, 1964, syn.nov.). The following revised or new combinations are proposed: Rhodochlanis Klimaszewski, 1967, stat. rev. (from Rhombaphalara); Rhodochlanis halocnemi (Loginova, 1964), comb.nov. (from Rhombaphalara); Rhodochlanis halostachidis (Loginova, 1970), comb.nov. (from Rhombaphalara); Rhodochlanis insolita (Burckhardt & Mifsud, 1998), comb.nov. (from Rhombaphalara). Identification keys are provided for adults of the world genera of Aphalarinae and for adults and larvae of the known Limataphalara species. Keywords. Psyllids, Aphalarinae, taxonomy, new species, phylogeny, host plants, Neotropical Region

Introduction Psyllids or jumping plant-lice are phloem-feeding Sternorrhyncha resembling minute cicadas. They are generally highly host specific on woody dicotyledonous plants and display their largest species diversity in the tropics (BURCKHARDT 2005, HOLLIS 2004). The subfamily Aphalarinae (Aphalaridae) is atypical in this respect as two thirds of its constituent genera occur in North temperate biomes and the two largest genera, Aphalara Foerster, 1848, and Craspedolepta Enderlein, 1921, are associated with herbaceous plants of the Asteraceae, Polygonaceae and several other families (Table 1). Aphalara and Craspedolepta are predominantly Holarctic in distribution; Crastina

35 D. BURCKHARDT & D. L. QUEIROZ

Loginova, 1964, and Colposcenia Enderlein, 1929, occur in the Palaearctic, Oriental and Afrotropical Regions; Brachystetha Loginova, 1964, Caillardia de Bergevin, 1931, Epheloscyta Loginova, 1976, Eumetoecus Loginova, 1961, Eurotica Loginova, Rhodochlanis Loginova, 1964, and Xenaphalara Loginova, 1961, are restricted to the Palaearctic; the monotypic genera Hodkinsonia Burckhardt et al., 2004, Lanthanaphalara Tuthill, 1959, Limataphalara Hodkinson, 1992, and Neaphalara Brown & Hodkinson, 1988, are endemic to the Neotropical Region; and Gyropsylla Brèthes, 1921, occurs in the Neotropical, Nearctic and Oriental Regions.

Table 1. Host plants of aphalarine genera.

Psyllid genus Host genus Aphalara Polygonaceae (Caryophyllales) and other families; herbaceous plants Brachystetha Nitraria (Nitrariaceae, Sapindales) Caillardia Anabasis, Haloxylon (Amaranthaceae, Caryophyllales) Colposcenia Tamarix (Tamaricaceae, Caryophyllales) Craspedolepta Asteraceae (Asterales) and other families; herbaceous plants Crastina Myricaria, Tamarix (Tamaricaceae, Caryophyllales) Epheloscyta Kalopanax (Araliaceae, Apiales) Eumetoecus Kochia (Amaranthaceae, Caryophyllales) Eurotica Eurotia (Amaranthaceae, Caryophyllales) Hodkinsonia Aralia (Araliaceae, Apiales) Gyropsylla Ilex (Aquifoliaceae, Aquifoliales) Lanthanaphalara Dunalia (Solanaceae, Solanales) Limataphalara Nectandra (Lauraceae, Laurales) Neaphalara Araliaceae (Apiales) Rhodochlanis Amaranthaceae (Caryophyllales) Xenaphalara Ceratocarpus (Amaranthaceae, Caryophyllales)

Based on adult characters of the Palaearctic taxa LOGINOVA (1964a, b) subdivided the Aphalarinae into the four tribes Aphalarini, Caillardiini, Colposceniini and Xenaphalarini. WHITE & HODKINSON (1985), in their classification of the world fauna, adopted to a large extent Loginova’s classification. They analysed adult and larval characters but included only six genera referred to this subfamily. They added the tribes Gyropsyllini for Gyropsylla and transferred the Phytolymini erected by BEKKER- MIGDISOVA (1973) for Phytolyma Scott, 1882, from the Anomalopsyllinae (Aphalaridae) to the Aphalarinae. The Phytolymini was later moved to the Homotomidae by OUVRARD (2002). KLIMASZEWSKI (1983, 1987) used a phenetic approach to analyse the Palaearctic taxa. He split Craspedolepta s. l. into several genera and subgenera and proposed a classification of six tribes which resembled that of WHITE & HODKINSON (1985) but differed in the erection of the Rhombaphalarini for three genera previously assigned to the Caillardiini. Klimaszewski’s classification was discussed by BURCKHARDT & LAUTERER (1997) who pointed out methodological shortcomings and mistakes in the

36 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara observation of some characters. KLIMASZEWSKI (2001) further developed his previous classifications and split off the Craspedoleptini from the Aphalarini. BURCKHARDT & OUVRARD (2012) recognised in their revised psyllid classification the extant tribes Aphalarini, Caillardiini, Colposceniini, Gyropsyllini and Xenaphalarini, suggesting that the subfamily in its present definition is likely to be monophyletic but that the constituent tribes may not be. In addition to the recent taxa, several fossil Aphalarinae have been described which are referred to the extinct tribe Paleopsylloidini (OUVRARD et al. 2013). The fossil tribe is not discussed here. In summary, none of the previous classifications of Aphalarinae used cladistic methodology nor included all genera in their assessments thus making them difficult to test. HODKINSON (1980, 1989, 1992) suggested that the majority of Aphalarinae is of Palaearctic origin, and that Aphalara and Craspedolepta migrated to North America across a Beringia connection. The Neotropical taxa, however, represent relict forms of a much larger fauna. Hodkinson (1992) pointed out that the small or monotypic Neotropical genera appear taxonomically isolated from each other but that the host plant, i.e. Nectandra (Lauraceae), could represent a shared character between related taxa. HOLLIS & MARTIN (1997) presented an overview of psyllids associated with Lauraceae and listed the following New World Aphalarinae from Nectandra: Gyropsylla cannela (Crawford, 1925) from ?Nectandra sp. (Brazil), Limataphalara brevicephala Hodkinson, 1992, from N. coriacea (Belize, USA: Florida), and a new species of an unidentified genus from N. membranacea (Costa Rica). During recent field work in Brazil we found another two undescribed species similar to the last one, also associated with Nectandra. Here we describe the new taxa from Nectandra, analyse their phylogenetic relationships within the Aphalarinae, test previous classifications of Aphalarinae and discuss host plant and biogeographic relationships.

Material and methods Material was examined or is listed from following institutions: LEEF ...... Laboratório Entomológico, Embrapa Florestas, Colombo, PR, Brazil MHNG ...... Muséum d’histoire naturelle, Genève, Switzerland MZSP ...... Museu de Zoologia, Universidade de São Paulo, SP, Brazil NHMB ...... Naturhistorisches Museum, Basel, Switzerland BMNH ...... Natural History Museum, London, UK USNM ...... United States National Museum collections, Beltsville, MD, USA

Morphological terminology follows HOLLIS (1976, 2004), BROWN & HODKINSON (1988), OSSIANNILSSON (1992), and YANG et al. (2009). The morphological studies were made with a Leica MZ12 stereo microscope and a Leica DMLB (with phase contrast and Nomarski differential interference contrast) compound microscope. The drawings were made with a drawing tube from permanent mounts in Canada balsam of dissected specimens previously cleared in KOH and washed in H2O. Measurements were taken from slide mounted material and are given in millimeters (mm).

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A cladistic analysis was performed with NONA version 2.0 (GOLOBOFF 1999) and WINCLADA 1.00 (NIXON 2002). The following parameters were used: maximum trees to keep = 10 000; number of replications = 5; starting trees per replication = 5; random seed = 0; search strategy = multiple TBR (tree-bisection-reconnection) + TBR; unconstrained search. For constructing a consensus tree the command Nelsen (collapse + consensus) was chosen.

Phylogenetic analysis For analysing the phylogenetic relationships among extant aphalarine genera we performed a cladistic analysis using 16 adult characters, six of which concern the head and its appendages, two the thorax, one the forewing, five the hind legs and two the aedeagus (Tables 2 and 3). Most characters are binary except for four, which are treated as unordered. Tainarys Brèthes, 1920 (Aphalaridae, Rhinocolinae) was included in the analysis as outgroup. The analysis resulted in 3 most parsimonious trees (Fig. 1, consensus tree) of 31 steps with a consistency index = 0.67 and retention index = 0.83. The anteriorly displaced eyes and enlarged postorbital ridges (characters 1 and 2, Table 2) in Eurotica and Xenaphalara were used by LOGINOVA (1964a) to group them

Table 2. Characters and states used in the cladistic analysis (cf. Table 3, Fig. 1).

1. Eyes in normal position (0); displaced anteriad (1) 2. Post-orbital ridges (in longitudinal body axis) short (0); long (1) 3. Vertex length > half width (0); equal or < half width (1) 4. Frons large, triangular (0); small, very short, parallel-sided (1); small, triangular (2); narrow, long, ribbon-shaped (3) 5. Clypeus flattened, heart-shaped (0); produced, spherical or tubular (1) 6. Rhinarium on antennal segment 7 absent (0); present (1) 7. Propleurites divided by a vertical suture (0); by an oblique suture (1) 8. Mesosternum subrectangular, rounded laterally (0); narrowed in the middle, with rim antero-laterally (1); very short, transverse (2) 9. Pterostigma of forewing present (0); absent (1) 10. Metacoxa with area beneath meracanthus straight (0); bearing a swelling (1) 11. Metatibia length/femur length ratio < 1.4 (0); > 1.5 (1) 12. Long stout bristles on apex of metafemur and base of metatibia lacking (0); present (1) 13. Apical metatibial spurs moderately dense, short (0); very dense, long (1); hardly sclerotised (2) 14. Metatarsus moderately long (0); very short (1) 15. Apex of aedeagus simple, weakly inflated (0); simple, strongly inflated (1); strongly inflated, with anterior screrotised hook and posterior membranous lobe (2) 16. Joint of distal segment of aedeagus at base (0); set some way from base (1)

38 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara

Fig. 1. Cladogram illustrating phylogenetic and host plant relationships of Aphalarinae. Tainarys (Rhinocolinae) was chosen as outgroup. Characters (numbers above dots) and character states are listed in Table 2, the character matrix in Table 3. Black dots represent synapomorphies, white dots homoplasies.

together in the Xenaphalarini. This group is also recovered in our analysis. KLIMASZEWSKI (1987) and LOGINOVA (1964a) split the subfamily into two groups based on the relative vertex length (character 3), short in the Caillardiini sensu Loginova and long in the other tribes. LOGINOVA (1964a) did not mention which state she thought was derived. KLIMASZEWSKI (1987) suggested, with doubts, that the short vertex is derived but he did not include the character in his cladogram. The Caillardiini sensu Loginova is also monophyletic in our analysis. A previously not considered character concerns the frons (character 4). In Tainarys (outgroup), Brachystetha and Colposcenia the frons is large and triangular suggesting that this is the plesiomorphic state. In Caillardia and Crastina the frons is very short and parallel-sided, in Eumetoecus and Rhodochlanis small and triangular, and in the remaining genera narrow, long and ribbon-shaped. The last two states define a monophyletic group each. The former corresponds to the Rhombaphalarini sensu KLIMASZEWSKI (1987), the latter to the tribes Aphalarini, Gyropsyllini and

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Table 3. Matrix for Aphalarinae (cf. Table 2, Fig. 1).

Tainarys 0000000000000000 Aphalara 0003110110000020 Brachystetha 0000000010000010 Caillardia 0011100000000010 Colposcenia 0000011000000020 Craspedolepta 0003110110000020 Crastina 0001110000000010 Epheloscyta 0003110100000010 Eumetoecus 0012100200000000 Eurotica 1103110110002000 Hodkinsonia 0003110100000010 Gyropsylla 0003111101111010 Lanthanaphalara 00031101011110?0 Limataphalara brevicephala 0013110101011101 Limataphalara eucosma 0013110101111101 Limataphalara hollisi 0013110101111111 Limataphalara lautereri 0013110101111111 Neaphalara 0003110100000010 Rhodochlanis 0012110200000010 Xenaphalara 1103110110002000

Xenaphalarini sensu WHITE & HODKINSON (1985). The clypeus (character 5) can be flattened as in Tainarys (outgroup), Brachystetha and Colposcenia, spherical or tubular. KLIMASZEWSKI (1987) used the clypeus in his analysis. The four states he distinguished are not clear and he may have miscoded some taxa. For example, Aphalara and Brachystetha are said to have a rounded forward directed clypeus, which is true for Aphalara but not Brachystetha where it is flattened. Most aphalarines have a single subapical rhinarium on each of segments 4–9 (character 6) but in Brachystetha, Caillardia and Eumetoecus those on segments 7 and sometimes 4 are reduced. KLIMASZEWSKI (1987) suggested that the presence of six rhinaria is primitive within Aphalarinae and that reductions occurred several times independently. This is confirmed by our study. The propleurites (character 7) are divided by a vertical suture in all genera except for Colposcenia and Gyropsylla where the suture is oblique. According to LOGINOVA (1964a) and WHITE & HODKINSON (1985) the suture is diagonal in Crastina but this is not the case. KLIMASZEWSKI (1987) suggested that this character is of little use to elucidate relationships in Aphalarinae which is confirmed by our analysis. The mesosternum (character 8) is subrectangular and rounded laterally in its plesiomorphic state. In Eumetoecus and Rhodochlanis it is very short and transverse. In the Epheloscyta–Limataphalara-clade it is narrowed in the middle with an antero-lateral rim. Several forewing characters have been used by previous authors in their analyses. KLIMASZEWSKI (1987) defined the Caillardiini sensu Loginova by forewings which ‘are thickened and half-cuticular’. This character is difficult to define and variable within some genera, and we consider it unsuitable for phylogenetic purposes. Similarly the

40 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara presence or absence of a ‘nodal-line’, a character which was used by previous authors, is sometimes difficult to determine. The suggested presence in Crastina by LOGINOVA (1964a) and WHITE & HODKINSON (1985) could not be substantiated. The same goes for the lack of a costal break in Aphalara and Craspedolepta mentioned by WHITE & HODKINSON (1985) which is erroneous. The presence/absence of a pterostigma (character 9) is equally problematical. It is absent in Aphalara, Brachystetha and Craspedolepta but present in the other genera though very narrow in some cases. In Gyropsylla, e.g. it is short and broad in G. cannela (Crawford, 1925) but long and very narrow in G. spegazziniana (Lizer, 1919). The clade Gyropsylla–Limataphalara is characterised by the metacoxa with the area beneath the meracanthus bearing a swelling (character 10). HODKINSON (1992) suggested that also Neaphalara bears this large swelling but this is not the case. The clade Gyropsylla–Limataphalara is further characterised by very long metafemora (character 11), the presence of long stout bristles on the apex of the metafemur and base of metatibia (character 12), and very dense, long apical metatibial spurs (character 13). In Eurotica and Xenaphalara the apical metatibial spurs are hardly sclerotised. Autaphomorphic for Limataphalara is the very short metatarsus (character 14). KLIMASZEWSKI (1987) used the shape of the apex of the distal segment of the aedeagus which we use here in a slightly simplified way (character 15). The analysis suggests that this character shows a lot of homoplasy. Autaphomorphic for Limataphalara is the joint of the distal segment of aedeagus which is set some way from base (character 16). No larval characters could be found suitable for the cladistic analysis mostly due to insufficient material.

Key to world genera of Aphalarinae (adults) 1 Metacoxa with area beneath meracanthus strongly swollen...... 2 – Metacoxa with area beneath meracanthus almost straight...... 4 2 Clypeus tubular, extended perpendicularly to lower head surface, with shoulder-like constriction medially. On Ilex (Aquifoliaceae)...... Gyropsylla – Clypeus short, subspherical...... 3 3 Vertex, along midline, longer than half its width, bearing a large anterior lobe on either half. On Dunalia (Solanaceae)...... Lanthanaphalara – Vertex, along midline, shorter than half its width, smoothly passing into genae anteriorly not bearing anterior lobes. On Nectandra (Lauraceae)...... Limataphalara 4 Eyes displaced anteriorly. Postorbital ridges large. Apical metatibial spurs light-coloured, hardly sclerotised...... 5 – Posterior eye margin almost level with posterior head margin. Postorbital ridges narrow. Apical metatibial spurs dark brown or almost black, strongly sclerotised...... 6

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5 Clypeus acute apically. Forewing with long, slender pterostigma, vein Rs sinuous relative to fore margin. On Eurotia (Amaranthaceae)...... Eurotica – Clypeus rounded apically. Forewing without pterostigma, vein Rs convex relative to fore margin. On Ceratocarpus (Amaranthaceae)...... Xenaphalara 6 Frons small, at most as long as longitudinal diameter of frontal ocellus, broad, parallel-sided...... 7 – Frons larger, more than twice as long as longitudinal diameter of frontal ocellus...... 8 7 Head inclined almost 90° from longitudinal body axis. On Anabasis and Haloxylon (Amaranthaceae)...... Caillardia – Head inclined about 45° from longitudinal body axis. On Myricaria and Tamarix (Tamaricaceae)...... Crastina 8 Frons widening towards base, more or less triangular, short or long...... 9 – Frons narrowly ribbon-shaped, often narrowed towards base, long...... 12 9 Clypeus adpressed, flattened. Vertex more than half as long as wide, ending in flat lobes anteriorly...... 10 – Clypeus sticking out, subglobular or tubular. Vertex less than half as long as wide, rounded to genae anteriorly...... 11 10 Propleurites with subequal proepisternum and epimeron, divided by perpendicular suture. Forewing lacking pterostigma. On Nitraria (Nitrariaceae)...... Brachystetha – Propleurites with small proepisternum and large epimeron, divided by diagonal suture. Forewing bearing distinct pterostigma. On Tamarix (Tamaricaceae)...... Colposcenia 11 Preocular sclerite large, posteriorly reaching to base of vertex. A subapical rhinarium present on each of antennal segments 4, 6, 8 and 9. On Kochia (Amaranthaceae)...... Eumetoecus – Preocular sclerite small, posteriorly reaching at most to the middle of vertex. A subapical rhinarium present on each of antennal segments 4–9. On various Amaranthaceae...... Rhodochlanis (= Rhombaphalara syn.nov.) 12 Forewing lacking pterostigma...... 13 – Forewing with membranous pterostigma...... 14 13 Clypeus tubular, often very long. Vertex angular anteriorly, separated from genae by narrow groove. On herbaceous Polygonaceae and other families...... Aphalara

42 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara

– Clypeus spherical. Vertex usually rounded anteriorly, passing smoothly into genae. On herbaceous Asteraceae and other families...... Craspedolepta 14 Pterostigma long, ending at about apical fifth of vein Rs. Palaearctic. On Kalopanax (Araliaceae)...... Epheloscyta – Pterostigma short, ending at about the middle or apical third of vein Rs. Neotropical...... 15 15 Genae with distinct tubercle on lower head surface between antennal insertion and eye. Clypeus short, in profile hardly visible. On Aralia (Araliaceae)...... Hodkinsonia – Genae lacking tubercle on lower head surface between antennal insertion and eye. Clypeus long, tubular, in profile clearly visible. On Araliaceae...... Neaphalara

Taxonomy Limataphalara Hodkinson, 1992 Limataphalara Hodkinson, 1992: 84. Type species Limataphalara brevicephala Hodkinson, 1992, by original designation and monotypy. Diagnosis. Adult. Head (Fig. 2) only little inclined from longitudinal body axis, moderately to very short. Vertex trapezoidal, two foveae distinct, coronal suture fully developed; anteriorly smoothly passing into genae; lateral ocelli situated adjacent to the eye. Frons forming narrow, elongate sclerite. Eyes hemispherical. Preocular sclerite developed at anterior half of eye, small or large. Antenna (Fig. 3) 10-segmented, 1.0–1.2 times as long as head width, segment 3 the longest, with a single large rhinarium on each of segments 4 to 9; terminal setae on segment 10 longer than segment, pointed. Clypeus small, pear-shaped, not extended anteriorly. Thorax weakly arched dorsally, with short, collar-like pronotum. Propleurites subrectangular, with subequal proepisternum and proepimeron, suture vertical with two dorsal branches. Mesoscutellum and metascutellum raised, knob-like. Mesosternum subrectangular, narrowed in the middle, with antero-lateral rim. Forewing (Figs 5–7) oblong-oval, with pterostigma and costal break, anal break vein adjacent to apex of vein Culb; vein R and M+Cu subequal, vein Rs almost straight, cell m1 smaller than cu1 which is large; marginal vein and apices of veins

Rs, M1+2, M3+4 and Cu1a conspicuously lighter than rest of veins and membrane; wing membrane semitransparent, veins concolorous or darker, particularly apically; surface spinules (Figs 8–10) covering all cells, leaving spinule-free stripes along veins, fine densely spaced, sometimes forming irregular transverse rows or rings. Hindwing membraneous, slightly shorter than forewing; furcation of vein R+M+Cu into R, M and Cu indistinct; costal setae grouped. Hind leg (Fig. 4) with metacoxa bearing a short, stout and apically rounded meracanthus and a rounded swelling between the meracanthus and the trochanther’s insertion; metafemur apically and sometimes metatibia basally with stout long bristles; metatibia 1.2–1.9 times as long as metafemur, lacking genual spine,

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Figs 2–10. Morphological details of Limataphalara spp. 2–4, 6, 9 – L. hollisi sp.nov. 5, 8 – L. eucosma sp.nov. 7, 10 – L. lautereri sp.nov. 2 – Head; 3 – antenna; 4 – hind leg; 5–7 – forewing; 8–10 – surface spinules in the apical third of cell m1. Scale for Figs 2–4: 0.2 mm; Figs 5–7: 0.2 mm; Figs 8–10: 0.05 mm.

44 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara

Figs 11–16. Terminalia of Limataphalara spp. 11, 14 – L. eucosma sp.nov. 12, 15 – L. hollisi sp.nov. 13, 16 – L. lautereri sp.nov. 11–13 – Male terminalia, lateral view; 14–16 – female terminalia, lateral view. Scale for Figs 11–13: 0.1 mm; Figs 14–16: 0.1 mm.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 45 D. BURCKHARDT & D. L. QUEIROZ with 7–10 strongly sclerotised apical spurs forming an open crown. Metabasitarsus very short, bearing two strongly sclerotised lateral spurs. Male proctiger (Figs 11–13) with moderately long, sinuous posterior processes usually bearing an inner, weakly sclerotised hook-like process; subgenital plate subglobular or elongate. Paramere (Figs 17–19) elongate, with a claw or thumb-like strongly sclerotised inner process situated between the basal and apical thirds of the anterior margin. Distal portion of aedeagus (Figs 20–22) articulated some way from base, bearing small or large subapical hook and long sclerotized end tube of ductus ejaculatorius with saw-shaped or denticulate dorsal margin. Female terminalia (Figs 14–16) cuneate; proctiger, in profile, arched distal to circumanal ring which consists of two rows of pores, the outer longitudinal to margin, the inner ones diagonal. Subgenital plate ranging from about half as long to almost as long as proctiger, ventral margin weakly curved or angled, pointed or incised apically. Fifth instar larva (Figs 26, 27, 29–31). Body oval, strongly flattened. Dorsal surface and margin densely covered in elongate tubercles bearing a lanceolate seta each; marginal tubercles longer than dorsal ones. Antenna 1- or indistinctly 2-segmented, covered in tubercles each bearing a single normal or lanceolate seta; one rhinarium clearly visible in basal part corresponding to adult scape, flagellar rhinaria indistinct (perhaps partially reduced). Clypeus with a pair of moderately long simple setae. Forewing bud with humeral lobe extending anteriorly to the middle of the eye. Thoracic tergites large. Caudal plate pointed posteriorly. Anus ventral, at some distance from the abdominal hind margin; circumanal ring small, lens-shaped, outer ring consisting of a single row of elongate pores. Tarsal arolium trapezoidal, shorter than claws, with short unguitractor, lacking petiole. Egg (Figs 23–25, 28). Oval with short lateral pedicel, widest in basal third, narrowing towards apex which bears a long filament which is almost as long as egg. Host plants. Limataphalara spp. appear monophagous on species of Nectandra Rol. ex Rottb. (Lauraceae). Distribution. Neotropical from the USA (Florida) in the north to Brazil (Paraná) in the south reflecting the distribution of its host genus Nectandra (ROHWER 1993).

Key to Limataphalara species: adults 1 Head, pro and mesothorax, legs and marginal half of forewing cell c+sc dark brown or almost black, rest of body and forewings amber- coloured. Metatibia/metafemur length ratio < 1.4. Belize, USA (Florida). On N. coriacea...... L. brevicephala Hodkinson – Body coloration different; vertex yellowish, ochreous or light brown; marginal area of forewing cell c+sc not significantly darker than remainder of wing (Figs 5–7). Metatibia/metafemur length ratio > 1.5...... 2

46 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara

2 Thorax dark brown or black dorsally with lighter longitudinal stripes on mesopraescutum and scutum and white spots. Body dimensions large: head width > 0.8 mm, forewing length > 2.3 mm. Paramere relatively massive with dense thick long setae along dorsal margin, and thick short claw-like process in the middle of the anterior margin (Figs 11, 17). Distal portion of aedeagus slender, styliform with small apical hook and serrate sclerotised end tube of ductus ejaculatorius (Fig. 20). Female subgenital plate pointed apically (Fig. 14). Brazil (Paraná). On N. lanceolata...... L. eucosma sp.nov. – Thorax ochreous or yellow dorsally with more or less expanded, dark brown or black median longitudinal stripe or band. Body dimensions smaller: head width < 0.8 mm, forewing length < 2.3 mm. Paramere relatively slender with sparse fine long setae along dorsal margin, and slender, long thumb-like process in the middle of the anterior margin (Figs 12, 13, 18, 19). Distal portion of aedeagus club-shaped with large apical hook and sclerotised end tube of ductus ejaculatorius bearing a single denticle (Figs 21, 22). Female subgenital plate incised apically Figs 15, 16)...... 3 3 Body dimensions larger: head width > 0.6 mm, forewing length > 1.9 mm. Paramere and distal portion of aedeagus as in Figs 12, 18. Female subgenital plate short: proctiger/subgenital plate ratio = 2.0; apical incision on subgenital plate long and wide. Costa Rica. On N. membranacea...... L. hollisi sp.nov. – Body dimensions smaller: head width < 0.6 mm, forewing length < 1.9 mm. Paramere and distal portion of aedeagus as in Figs 13, 19. Female subgenital plate long: proctiger/subgenital plate ratio = 1.6; apical incision on subgenital plate short and narrow. Brazil (Mato Grosso, Pará). On N. cuspidata...... L. lautereri sp.nov.

Key to Limataphalara species: fifth instar larvae [larvae of L. brevicephala and L. hollisi unknown] 1 Forewing pad long, antenna length/forewing pad length ratio < 0.5. Setae on antenna shorter than diameter of antenna; lanceolate setae on margins of head, wing pads and caudal plate shorter than tubercles on which they are situated...... L. eucosma sp.nov. – Forewing pad short, antenna length/forewing pad length ratio > 0.5. Setae on antenna longer than diameter of antenna; lanceolate setae on margins of head, wing pads and caudal plate longer than tubercles on which they are situated...... L. lautereri sp.nov.

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List of species Limataphalara brevicephala Hodkinson, 1992

Limataphalara brevicephala Hodkinson, 1992: 86; holotype ♂, USA: Florida, Miami, 13.iv.1982, Nectandra coriacea (USNM; not examined).

Material examined. Belize: 1 ♂, Cayo, Chiquibul Forest, Las Cuevas, 21.iii.1995, Fog 10, Cedrela, D. Hollis et al. leg.; 1 ♀, same but iii–iv.1999, C. Minty leg. (MT) (BMNH, dry mounted).

Description. Adult. Described by HODKINSON (1992). Fifth instar larva. Unknown. Host plant. Nectandra coriacea (Sw.) Griseb. (Lauraceae). Distribution. USA (Florida) (HODKINSON 1992), Belize (HOLLIS & MARTIN 1997).

Limataphalara eucosma sp.nov. (Figs 5, 8, 11, 14, 17, 20, 23, 26, 28–31)

Type material. Holotype: ♂, Brazil: Paraná: Colombo, S25°19.078′ W49°09.116′, 930 m, 24.v.2013, Nectandra lanceolata, D. L. Queiroz leg. (MZSP, slide mounted). Paratypes: Brazil: Paraná: 1 ♂, 3 ♀♀, 3 larvae, 3 parasitised larval cases, same data as holotype. 1 ♂, 2 ♀♀, same but 17.vii.2013, S25°19.096′ W49°09.100′, D. L. Queiroz leg., #542. Rio Grande do Sul: 1 ♂, RS, Passo Fundo, 27.vi.2013, S28°13.195′ W52°24.714, 630 m, D. L. Queiroz leg. #517 (LEEF, MZSP, NHMB, slide mounted, and preserved in 70% ethanol). Description. Adult. Coloration. Ochreous with brown, black and white patches and dots. Vertex light brown, darker anteriorly, anterior margin white, anterior delimitation to genae and occiput black. Eyes grey, ocelli reddish. Lower head surface brown to black. Antenna yellowish, segments 1 and 2 brown, segments 4–8 with dark apices, segments 9 and 10 dark brown or black. Thorax dark brown or black dorsally with lighter longitudinal stripes on mesopraescutum and scutum and white spots as follows: six on pronotum, and two each on posterior margin of mesoprascutum, meso and metascutellum. Thorax laterally and ventrally straw-coloured with dark margins of sclerites. Forewing with yellowish or light brown, transparent membrane and basally yellow and otherwise brown to black veins, apices of veins Rs, M1+2, M3+4 and Cu1a white. Hindwing whitish, transparent. Legs yellowish to brown. Abdominal sclerites including terminalia ochreous to dark brown, intersegmental membrane yellow. Structure. Body dimensions large. Surface spinules of forewing (Fig. 8) arranged in irregular transverse rows. Terminalia as in Figs 11, 14, 17, 20. Male subgenital plate elongate, ventral margin, in profile, angular. Paramere relatively broad with a claw-like hook slightly above the middle of the fore margin, a sclerotised rim near apical margin and an inward directed sclerotised point near apex on the inner face; inner face in apical third of posterior half with tubercular microsculpture; dorsal margin with long, dense setae. Distal segment of aedeagus slender with small apical hook and moderately long, sinuous sclerotised end tube of the ductus ejaculatorius which is serrate on its dorsal side. Female subgenital plate long, pointed apically. Measurements and ratios in Table 4.

48 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara

Figs 17–22. Male genital organs of Limataphalara spp. 17, 20 – L. eucosma sp.nov. 18, 21 – L. hollisi sp.nov. 19, 22 – L. lautereri sp.nov. 17–19 – Paramere, inner face; 20–22 – distal portion of aedeagus. Scale: 0.05 mm.

Fifth instar larva. Coloration. Ochreous with dark brown spines. Eyes grey. Antenna yellow with dark tip. Structure. Lanceolate setae on antenna shorter than antennal diameter. Marginal tubercles longer than associated lancelate seta. Tubercles on thoracic and abdominal dorsum in average about twice as long as their base. Forewing pad long. Measurements and ratios in Table 5. Etymology. From Greek ευκοσµος = ornate, referring to the attractive adult colour pattern. Host plant. Nectandra lanceolata Nees & Mart. (Lauraceae). Distribution. Brazil (Paraná).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 49 D. BURCKHARDT & D. L. QUEIROZ

Table 4. Measurements (in mm) and ratios of adult Limataphalara spp. (n = 1 ♂, 1♀/ species).

head antenna forewing metatibia ♂proctiger paramere aedeagus ♀proctiger width length length length length length distal sgmt length (HW) (AL) (WL) (TL) (MP) (PL) length (FP) eucosma ♂ 0.80 0.80 2.30 0.70 0.18 0.25 0.25 eucosma ♀ 0.88 0.90 2.63 0.78 0.70 hollisi ♂ 0.65 0.75 2.00 0.55 0.15 0.20 0.25 hollisi ♀ 0.73 0.78 2.30 0.60 0.50 lautereri ♂ 0.55 0.60 1.53 0.48 0.13 0.18 0.20 lautereri ♀ 0.60 0.60 1.78 0.50 0.45

AL/ WL/ WL/ TL/ TL/ MP/ FP/ FP/ FP/ ♀ HW HW forewing HW metafemur HW HW circumanal subgenital breadth length ring length plate eucosma ♂ 1.00 2.88 2.30 0.88 1.87 0.22 eucosma ♀ 1.03 3.00 2.19 0.89 1.63 0.80 2.80 1.56 hollisi ♂ 1.15 3.08 2.29 0.85 1.83 0.23 hollisi ♀ 1.07 3.17 2.30 0.83 1.71 0.69 1.82 2.00 lautereri ♂ 1.09 2.77 2.18 0.86 1.73 0.23 lautereri ♀ 1.00 2.96 2.09 0.83 1.67 0.75 2.00 1.06

Limataphalara hollisi sp.nov. (Figs 2–4, 6, 9, 12, 15, 18, 21, 24)

Psyllidae Aphalarinae genus and species n., HOLLIS & MARTIN 1997: 471, table 1.

Type material. Holotype: ♂, Costa Rica: San José, 2–4 km East of San Jeronimo, 1500 m, 4.ii.1993, Nectandra membranacea, D. Hollis leg. (BMNH, dry mounted). Paratypes: Costa Rica: 1 ♂, 2 ♀♀, same data as holotype; 1 ♂, 1 ♀, same data as holotype but 26.i.–4.ii.1993 (BMNH, NHMB, dry and slide mounted). Description. Adult. Coloration. Yellow or straw-coloured dorsally, brown or dark brown ventrally. Antennal segments 1 and 2 ochreous or brown, 3–7 yellow, 4–8 dark brown apically, 9 and 10 almost black. Clypeus yellow. Meso- and metanotum with more or less expanded dark brown median longitudinal stripe. Forewing semi-transparent, with greyish membrane und concolorous veins except for apical region where veins are almost black, strongly contrasting with the membrane and apices of veins Rs, M1+2, M3+4 and Cu1a, which are white. Hindwing colourless. Femora and metacoxae dark brown. Abdominal sclerites dark brown, membranes yellow. Male proctiger ochreous. Female terminalia yellow. Younger specimens with less expanded dark colour. Structure. Body dimensions intermediate between those of L. eucosma and lautereri. Surface spinules of forewing (Fig. 9) arranged in irregular rings. Terminalia as in Figs 12, 15, 18, 21. Male subgenital plate subglobular, ventral margin, in profile, broadly rounded. Paramere relatively slender with long thumb-like process in apical third of the fore margin and sclerotised apical margin; inner face in apical third of posterior half lacking tubercular microsculpture; dorsal margin with moderately long, spaced setae. Distal

50 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara

Figs 23–31. Eggs and larvae of Limataphalara spp. 23, 26, 28–31 – L. eucosma sp.nov. 24 – L. hollisi sp.nov. 25, 27 – L. lautereri sp.nov. 23–25 – Eggs in female abdomen; 26, 27 – fifth instar larva (26 – left dorsal, right ventral view); 28 – egg on branchlet of Nectandra lanceolata; 29 – deformation induced on N. lanceolata; 30, 31 – larvae on branchlet of Nectandra lanceolata. Scale for Figs 23–25: 0.1 mm; Figs 26, 27: 0.2 mm.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 51 D. BURCKHARDT & D. L. QUEIROZ

Table 5. Measurements (in mm) and ratios of fifth instar larvae of Limataphalara spp. (n: L. eucosma = 1; L. lautereri = 2).

body antenna BL/body BB/ caudal AL/ caudal CPB/ circumanal length length breadth plate breadth forewing pad plate length ring (BL) (AL) (BB) (CPB) length / CPB breadth eucosma 1.53 0.38 1.05 1.61 0.40 4.00 6.55 lautereri 1.40–1.48 0.34–0.44 1.18–1.19 1.62–1.72 0.53–0.78 3.00–5.00 5.27–7.25

segment of aedeagus thick with large apical hook and moderately long, sinuous sclerotised end tube of the ductus ejaculatorius, which bears a tubercle on its dorsal side. Female subgenital plate short, with deep, wide incision apically. Measurements and ratios in Table 4. Fifth instar larva unknown. Etymology. Dedicated to David Hollis who collected the type material. Host plant. Nectandra membranacea (Sw.) Griseb. (Lauraceae). Distribution. Costa Rica.

Limataphalara lautereri sp.nov. (Figs 7, 10, 13, 16, 19, 22, 25, 27)

Type material. Holotype: ♂, Brazil: Pará: Belém, Embrapa campus, S1°24.9–26.6′ W48°25.3–26.6′, 20 m, 8–15.iv.2013, Nectandra cuspidata, edge of Amazonian forest, D. Burckhardt & D. L. Queiroz leg., #99(2) (MZSP, dry mounted). Paratypes: Brazil: Mato Grosso: 2 ♀♀, Tabaporã, Fazenda Crestani, S11°18.8/20.2′ W55°57.7/58.5′, 330–380 m, 6–8.xi.2012, transitional forest near river, forest edges, farm land, parc vegetation, D. Burckhardt & D. L. Queiroz leg., #62; 1 ♀, same but Tabaporã, S11°20.0′ W55°50.5′, 430 m, 8.xi.2012, forest edge along unpaved road, D. Burckhardt & D. L. Queiroz leg., #63. 1 ♂, 4 larvae, 1 skin, Sinop, 28.viii.2013, S11°52.250′ W55°35.747′, 350 m, D. L. Queiroz leg., #566; 2 ♂♂, same but 29.viii.2013, S11°52.967′ W55°38.420′, 320 m, #569; 2 ♀♀, same but 30.viii.2013, S11°52.252′ W55°35.746′, 360 m, #571; 2 ♂♂, 2 ♀♀, same but S11°52.164′ W55°35.804′, #572; 2 ♂♂, 4 ♀♀, 19 imm, 14 skins, Ponta Pora, 12.ix.2013, S21°59.940′ W55°33.814′, 585 m, Fazenda Mariana, D. L. Queiroz leg., #576; 1 ♀, Bonito, 19.ix.2013, S21°10.282′ W56°26.457′, 290 m, Hotel Cabanas ground, D. L. Queiroz leg. #585; 1 ♂, 1 ♀ same but S21°10.345′ W56°26.533′, 320 m, #586. Pará: 119 ♂♂, 92 ♀♀, 2 larvae, 3 parasitised larval cases, same data as holotype; 8 ♂♂, 14 ♀♀, 4 larvae, Pará, Belém, Jardim Botânico Bosque Rodrigues Alves, S1°25.8′ W48°27.2′, 30 m, 14.iv.2013, Nectandra cuspidata, edge of Amazonian forest; sweeping vegetation, D. Burckhardt & D. L. Queiroz leg., #103(2) (LEEF, MHNG, MZSP, HNMB, BMNH, USNM; dry and slide mounted and preserved in 70% and 100% ethanol). Description. Adult. Coloration. Ochreous. Head black ventrally. Antennal segments 1 and 2 brown, 3 entirely yellow, 4–8 yellow basally and brown apically, 9 and 10 almost black. Pronotum yellow; thorax with black longitudinal median stripe extending from mesoscutum to metascutellum. Forewing semi-transparent, ochreous, veins only slightly darker than membrane, apices of veins Rs, M1+2, M3+4 and Cu1a white. Hindwing colourless, transparent. Abdominal sternites in male dark brown or black. Structure. Body dimensions small. Surface spinules of forewing (Fig. 10) irregularly, densely arranged. Terminalia as in Figs 13, 16, 19, 22. Male subgenital plate

52 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara subglobular, ventral margin, in profile, weakly curved. Paramere relatively slender with short thumb-like process in apical third of the fore margin and sclerotised apical margin; inner face in apical third of posterior half with tubercular microsculpture; dorsal margin with moderatly long, spaced setae. Distal segment of aedeagus moderately thick with large apical hook and moderately long, sinuous sclerotised end tube of the ductus ejaculatorius which bears a tubercle on its dorsal side. Female subgenital plate short, with shallow, narrow incision apically. Measurements and ratios in Table 4. Fifth instar larva. Coloration. Yellowish, spines slightly darker. Structure. Lanceolate setae on antenna longer than antennal diameter. Marginal tubercles much shorter than associated lancelate seta. Tubercles on thoracic and abdominal dorsum in average about as long as their base. Forewing pad short. Measurements and ratios in Table 5. Etymology. Dedicated to Pavel Lauterer. Host plant. Nectandra cuspidata Nees & Mart. (Lauraceae). Distribution. Brazil (Mato Grosso, Pará).

Discussion and conclusions Apart from the monotypic Gyropsyllini, following previously defined groupings appear also in our cladistic analysis (Fig. 1, Tables 2 and 3): the Xenaphalarini, the Caillardiini sensu LOGINOVA (1964a) and the Rhombaphalarini of previous authors (KLIMASZEWSKI 1987, 2001, LOGINOVA 1964a, WHITE & HODKINSON 1985). Otherwise our analysis is considerably different suggesting the tribes Aphalarini, Craspedoleptini and Stigmaphalarini of previous classifications are paraphyletic or polyphyletic. This is mostly due to the position of Brachystetha and Crastina. The former differs from Aphalara and close relatives in the large, triangular frons, the flattened clypeus and the subrectangular, laterally rounded mesosternum. The latter differs from Colposcenia in the small, paralell-sided frons, the subglobular clypeus and the vertical propleural suture. The relationships of Brachystetha and Colposcenia with respect to the remaining genera are not resolved in our analysis. The clades Crastina–Limataphalara, Eumetoecus+ Rhodochlanis, Epheloscyta–Limataphalara, Eurotica+Xenaphalara and Gyropsylla– Limataphalara are well supported. There are no characters, apart from the common host family Araliaceae, to group the two Central American genera Hodkinsonia and Neaphalara as well as the eastern Asian Epheloscyta together. The two Central American genera do not, however, form a distinct group with the other Neotropical members of the subfamily (Gyropsylla, Lanthanaphalara and Limataphalara). Applying a formal tribal classification would result in at least two monotypic tribes, depending on where the level is set, which would be of no practical use. We therefore synonymise all extant tribes: Aphalarinae Löw, 1879: 606 (= Caillardiini Loginova, 1964: 447, syn.nov.; Coelocarinae Li, 2011: 351, syn.nov.; Colposceniini Bekker-Migdisova, 1973: 109, syn.nov.; Craspedoleptini Klimaszewski, 2001: 196, syn.nov.; Eumetoecini

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 53 D. BURCKHARDT & D. L. QUEIROZ

Li, 2011: 356, syn.nov.; Gyropsyllini White & Hodkinson, 1985: 270, syn.nov.; Rhombaphalarini Klimaszewski, 1987: 228, syn.nov.; Stigmaphalarini Vondráèek, 1957: 140, nomen nudum, syn.nov.; Xenaphalarini Loginova, 1964: 447, syn.nov.). CONCI & TAMANINI (1984) and BURCKHARDT (1989) discussed Rhodochlanis and Rhombaphalara and suggested that they may be congeneric. They did not, however, formally synonymise them awaiting an analysis of the phylogenetic relationships within the subfamily. In our analysis we found no morphological differences between the two and formally synonymise them here: Rhodochlanis Loginova, 1964 (= Rhombaphalara Loginova, 1964, type species Rhombaphalara halocnemi Loginova, 1964, by original designation and monotypy, syn.nov.). The following revised or new combinations are proposed: Rhodochlanis achaetae Klimaszewski, 1967, stat. rev. (from Rhombaphalara); Rhodochlanis halocnemi (Loginova, 1964), comb.nov. (from Rhombaphalara); Rhodochlanis halostachidis (Loginova, 1970), comb.nov. (from Rhombaphalara); Rhodochlanis insolita (Burckhardt & Mifsud, 1998), comb.nov. (from Rhombaphalara). HODKINSON (1980) suggested the presence of Aphalara and Craspedolepta in the Nearctic region may be due to immigration from the Palaearctic region across a Beringia connection. On the other hand Gyropsylla, Hodkinsonia and Neaphalara were regarded by HODKINSON (1989) as an old Neotropical group. A Palaearctic origin for Aphalara and Craspedolepta is in accordance with our cladistic analysis. The biogeographical relationships of the other three New World genera seem more complicated but without a fully resolved phylogeny little can be concluded at present. Gyropsylla has an interesting distribution that includes both the Neotropics and China. The phylogenetic relationships of the Araliaceae feeding genera Epheloscyta, Hodkinsonia and Neaphalara are unresolved. If these genera should prove be monophyletic as a group then this would be another case of a Neotropical–eastern Asian distribution. The host relationships within the Aphalarinae are visualised in Fig. 1. Five genera are associated with Amaranthaceae, three with Araliaceae, two with Tamaricaceae and the rest with a single plant family each. Neither the genera associated with Amaranthaceae nor those with Tamaricaceae form a monophyletic group, suggesting that the two plant families were colonised by psyllids more than once. As the phylogenetic relationships of the three psyllid genera associated with Araliaceae are not resolved nothing can be concluded. The host of Gyropsylla cannela has been reported as possibly Nectandra sp. This could not be substantiated, however, as we found a series of G. cannela, adults and larvae, on Ilex microdonta during recent field work in Brazil (Burckhardt & Queiroz, unpublished data). This host is more plausible as the other Gyropsylla species with known host plants all develop on Ilex.

Acknowledgements We dedicate this paper to our friend Pavel Lauterer whose outstanding research on psyllids significantly contributed to the knowledge on psyllid biology, faunistics and systematics. We thank Diana Percy for the loan of material as well as Ian D. Hodkinson and Igor Malenovský for useful comments on an earlier manuscript draft. Osmar S. Ribas

54 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Phylogenetic relationships within Aphalarinae and revision of Limataphalara kindly identified plant samples for which we are grateful. For the specimens from Brazil we received the collecting permits numbers SISBIO 11832-2 and MCTI 294/2013 as well as 002152/2012-6.

References

BEKKER-MIGDISOVA E. E. 1973: Sistema psillomorf (Psyllomorpha) i polozhenie gruppy v otryade ravnokrylykh (Homoptera). [On the systematics of the Psyllomorpha and the position of the group within the order Homoptera.] Pp. 90–117. In: NARCHUK E. P. (ed.): Doklady na dvadzat chetvertom ezheghodnom chtenii pamyati N. A. Kholodkovskogo 1-2 Aprelya 1971 g. Voprosy paleontologii nasekomykh. Nauka, Leningrad, 118 pp (in Russian). BROUN R. G. & HODKINSON I. D. 1988: Taxonomy and ecology of the jumping plant-lice of Panama (Homoptera: Psylloidea). Entomonograph. Volume 9. Leiden, New York, Kopenhagen, Köln: E. J. Brill, Scandinavian Science Press Ltd, 304 pp. BURCKHARDT D. 1989: Les psylles (Insecta, Homoptera, Psylloidae) de l’Algérie. Archives des Sciences, Genève 42: 367–424. BURCKHARDT D. 2005: Biology, ecology, and evolution of gall-inducing Psyllids (Hemiptera: Psylloidea). Pp. 143–157. In: RAMAN R., SCHAEFER C. W. & WITHERS T. M. (eds.): Biology, ecology, and evolution of gall- inducing arthropods. Science Publishers, Enfield – Plymouth, xxi + 817 pp. BURCKHARDT D. & LAUTERER P. 1997: Systematics and biology of the Aphalara exilis (Weber and Mohr) species assemblage (Hemiptera: Psyllidae). Entomologica Scandinavica 28: 271–305. BURCKHARDT D. & OUVRARD D. 2012: A revised classification of the jumping plant-lice (Hemiptera: Psylloidea). Zootaxa 3509: 1–34. CONCI C. & TAMANINI L. 1984: Rhodochlanis salicorniae Klim., nuovo per l’Italia, R. hodkinsoni n. sp., Puglia, da Suaeda vera, e considerazioni sul genere (Homoptera Psylloidea Aphalaridae). Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 125: 61–80. GOLOBOFF P. 1999: NONA (NO NAME). Version 2. Published by the author, Tucumán, Argentina. Computer program available from http://www.cladistics.com HODKINSON I. D. 1980: Present day distribution patterns of the holarctic Psylloidea (Homoptera: Insecta) with particular reference to the origin of the nearctic fauna. Journal of Biogeography 7: 127–146. HODKINSON I. D. 1989: The biogeography of the Neotropical jumping plant-lice (Insecta: Homoptera: Psylloidea). Journal of Biogeography 16: 203–217. HODKINSON I. D. 1992: A new genus of Psyllid (Homoptera: Psylloidea: Aphalaridae) from Florida. Florida Entomologist 75: 84–89. HOLLIS D. 1976: Jumping plant lice of the tribe Ciriacremini (Homoptera: Psylloidea) in the Ethiopian Region. Bulletin of the British Museum Natural History (Entomology), London 34: 3–83. HOLLIS D. 2004: Australian Psylloidea: jumping plantlice and lerp insects. Australian Biological Resources Study, Canberra, xvi + 216 pp. HOLLIS D. & MARTIN J. H. 1997: Jumping plantlice (Hemiptera: Psylloidea) attacking avocado pear trees, Persea americana, in the New World, with a review of Lauraceae-feeding amongst psylloids. Bulletin of Entomological Research 87: 471–480. KLIMASZEWSKI S. M. 1983: Revision of the Palaearctic species of the genus Craspedolepta Enderl. s. l. (Homoptera, Aphalardide). Polskie Pismo Entomologiczne 53: 3–29. KLIMASZEWSKI S. M. 1987: Relationships within the Aphalarinae Loew subfamily (Homoptera, Psylloidea). Polskie Pismo Entomologiczne 57: 211–234. KLIMASZEWSKI S. M. 2001: Jumping plant lice from the family Aphalaridae in Central Asian Province (Hemiptera: Sternorrhyncha: Psyllolidea). Annals of the Upper Silesian Museum (Entomology) 10–11: 179–201. LOGINOVA M. M. 1964a: 2. Podotryad Psyllinea – Psyllidy, ili listobloshki. [Suborder Psyllinea – Psyllids or jumping plant-lice.] Pp. 437–482. In: BEI-BIENKO G. I. (ed.): Opredelitel nasekomykh evropeiskoi chasti SSSR. [Key to the insects of the European part of the USSR]. Nauka, Moskva – Leningrad, 935 pp (in Russian).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 55 D. BURCKHARDT & D. L. QUEIROZ

LOGINOVA M. M. 1964b: Novye i maloizvestnye psillidy Kazakhstana. Zametki o sisteme i klassifikatsii Psylloidea (Homoptera). [New and little known psyllids of Kazakhstan. Notes on the system and classification of Psylloidea (Homoptera).] Trudy Zoologicheskogo Instituta Akademii nauk SSSR 34: 52–112 (in Russian). NIXON K. C. 2002: WinClada. Version 1.00.08. Published by the author, Ithaca, NY, USA. Computer program available from http://www.cladistics.com OSSIANNILSSON F. 1992: The Psylloidea (Homoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica. Volume 26. E. J. Brill, Leiden – New York – Köln, 347 pp. OUVRARD D. 2002: Systématique phylogénetique des Hemiptera Psylloidea: Morphologie comparée du thorax et structures secondaires de l’ARNr 18S. (Phylogenetic systematics of Hemiptera Psylloidea: Comparative morphology of the thorax and secondary structures of the 18S rRNA.) Bulletin de la Société Zoologique de France 127: 345–357. OUVRARD D., BURCKHARDT D. & GREENWALT D. 2013: The oldest jumping plant-louse (Insecta: Hemiptera: Sternorrhyncha) with comments on the classification and nomenclature of the Palaeogene Psylloidea. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2):: 21–33. ROHWER J. G. 1993: Lauraceae: Nectandra. Flora Neotropica, monograph 60: 1–332. WHITE I. M. & HODKINSON I. D. 1985: Nymphal taxonomy and systematics of Psylloidea (Homoptera). Bulletin of the British Museum (Natural History), Entomology series 50: 153–301. YANG M.-M., BURCKHARDT D. & FANG S.-J. 2009: Psylloidea of Taiwan. Volume I. Families Calophyidae, Carsidaridae, Homotomidae und , with overview and keys to families and genera of Taiwanese Psylloidea (Insecta: Hemiptera). National Chung Hsing University, Taichung, 96 pp.

56 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 57–95, 2013

Revealing the identity of some early described European Cixiidae (Hemiptera: Auchenorrhyncha) – a case of ‘forensic’ taxonomy; two new combinations and a name change for Reptalus panzeri in Britain

MICHAEL D. WEBB1, ALEX J. RAMSAY2 & VALÉRIE A. LEMAÎTRE1 1 Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD; e-mails: [email protected]; [email protected] 2 44 Sun Lane, Burley-in-Wharfedale, Ilkley, West Yorkshire LS29 7JB; e-mail: [email protected]

WEBB M. D., RAMSAY A. J. & LEMAÎTRE V. A. 2013: Revealing the identity of some early described European Cixiidae (Hemiptera: Auchenorrhyncha) – a case of ‘forensic’ taxonomy; two new combinations and a name change for Reptalus panzeri in Britain. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 57–95. – Despite recent work on the identity of European Cixiidae, some anomalies have been found to remain. For example, just prior to this study, while identifying a British Reptalus Emeljanov, 1971 specimen we found it to match R. panzeri (Löw, 1883) in the British handbook, but also matched R. quinquecostatus (Dufour, 1833) of continental European authors. Further investigations revealed that the syntypes of Dufour’s species matched neither of the above identifications but in fact matched the figures given by authors of R. melanochaetus (Fieber, 1876). These findings lead us to investigate how these and certain other early identifications of European Cixiidae had been made and how the British Reptalus species had remained misidentified for so long. During the course of the above work it was necessary to consult the historical literature relating to the above species and some of the more important articles are reproduced here, together with English translations for the first time. One of the studies consulted was FIEBER’s (1875) ‘The Cicadines of Europe’ Part 1. In this work, Fieber noted he had figured in colour all the type specimens that he could examine but these are not to be found in Fieber’s work. However, from an indication in the translator’s Preface the figures were discovered in the Muséum national d’Histoire naturelle, Paris, where they had remained unreported since their acquisition. As a result of the above work we argue for the retention of the current identity of Reptalus quinquecostatus and R. melanochaetus in continental Europe and change the name for the British species R. panzeri to R. quinquecostatus, together with ecological and distributional notes on the latter and a revised key to separate the British genera of Cixiidae. In addition, the identities of two junior synonyms of Pentastiridius leporinus (Linnaeus, 1761) (Flata pallens Germar, 1821 and F. pallida Herrich-Schäffer, 1835), are also considered and resulting from a study of Fieber’s figures, noted above, the male genitalia of Reptalus limbatus (Fieber, 1876) comb. nov., were found to be very similar to R. venosus (Rambur, 1840) comb. nov. Detailed figures of the former are given for the first time. From FIEBER’s (1875) mention of the word ‘type’, noted in the current work, we take the opportunity to document the history of the ‘type concept’ and the earliest usage of the word ‘type’ (see Appendix 13).

Keywords. Fulgoromorpha, nomenclature, taxonomy, type concept, key to British Cixiidae

“Many of the [cixiid] species are exceedingly like each other, and this has led to great confusion. Authors have not known what to do with them, and so they have been separated by one, put together by another, and finally mixed up almost indiscriminately.” SCOTT (1870)

57 M. D. WEBB ET AL.

Introduction The planthopper family Cixiidae comprises approximately 146 genera and 2000 species worldwide (HOLZINGER et al. 2002); thus the family is one of the largest in the Fulgoromorpha (Hemiptera: Auchenorrhyncha). Some members of the family are notable for their subterranean nymphs and the wax they and the adults produce (Plate 1). Also, as mainly phloem feeders, the sap-sucking habit of the group makes them potential vectors of phloem-inhabiting plant pathogens and indeed several European species are known virus vectors while some species of Reptalus, treated here, are potential vectors of Phytoplasmas (BERTIN et al, 2010). Although found mainly in the tropics, the first species descriptions were mainly of European species. However, these early descriptions were so imprecise that each could ‘fit’ more than one species, although in some cases locality data may prove helpful (as for example with the identity of the earliest described European cixiids, Pentastiridius leporinus (Linnaeus, 1761) (see Results). Therefore, the ‘true’ identity of early described species can be unclear when their original descriptions lack certain diagnostic features that are in current use. This problem is compounded when type specimens are either unknown or are difficult to identify due to the absence of data in the original description and/or on the specimens. Also, in the absence of any early type concept any extant type material may also lack type identification labels. Few nineteenth century authors in the Hemiptera: Auchenorrhyncha referred to borrowed type material when identifying species. Such trafficking of material might have been more difficult than today, particularly as specimens were more frequently in private hands, prior to being placed in institutions. Nonetheless, for some major revisionary works, e.g. FIEBER (1875, 1876, 1878, 1879) and MELICHAR (1896), types were borrowed, referred to in the former case as either the ‘type’ or ‘original’ material (see below). For other authors, if comparative original (type) material was not at their disposal, only brief published original descriptions could be relied on for identifications. Nowadays, when confronted with a similar situation an entomologist usually has little choice than to follow historical perceptions and a consensus of opinion in the literature on the identity of a given species. This course of action results in stability, at least until a missing putative type is found that contradicts the accepted view. As with most Auchenorrhyncha, the identity of most cixiids is now based predominantly on characters of the male genitalia, however these characters were not generally used in early descriptions or were described but not figured, e.g. FIEBER (1876). It is unsurprising therefore, that this fact, together with unavailable type specimens and the large literature for some species (e.g., there are nearly 100 references to Pentastiridius leporinus from 1761 to 1929, in METCALF’s (1936) catalogue), has given rise to many misidentifications in the literature of this group. The problem of misidentifications in the literature was recognised early when SCOTT (1870) commentated on the confusion of British Cixiidae (see above quote on title page). Scott went on to say that the male styles serve as a “great guide” for identification, but although this proved to be the case, such characters only serve to distinguish species, not to apply the name, and in this instance Scott, like other authors (before and after), got some of the names wrong (see below).

58 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Identity of some early described European Cixiidae

The separation of European Cixiidae genera has frequently been based on the number of keels on the mesononotum. For example, LE QUESNE & PAYNE (1981), in their checklist of the British Auchenorrhyncha, recognise 12 Cixiidae species placed in two genera, Latreille, 1804 and Oliarus Stål, 1862, based on the presence of three or five ‘scutellum’ keels, respectively. These British species are currently placed in five genera of which only two (Pentastiridius Kirschbaum, 1868 and Reptalus Emeljanov, 1971) have five keels. Each of these genera comprises a single British species, but the identity of these two species was confused in the early literature in both Britain and Continental Europe, a situation that we have found still exists today. During routine identification work on a British Reptalus specimen (RAMSAY 2009) it was found to match the figures of R. panzeri (Löw, 1883) in the British Plate 1. Subterranean immature of Reptalus panzeri handbook (LE QUESNE 1960), but also (Löw, 1833). Image provided by Gernot Kunz. matched the figures of R. quinquecostatus (Dufour, 1833) given by continental authors (e.g. HOLZINGER et al. 2003). In order to check which identification was correct we borrowed the syntypes of Dufour’s species housed in the Muséum national d’Histoire naturelle, Paris. Much to our surprise we found that the types matched neither of the above identifications but in fact matched the figures given by authors of R. melanochaetus (Fieber, 1876)! These findings led to further investigations to establish how certain early identifications of European Cixiidae had been made and how the British Reptalus species had remained misidentified for so long. In the current study we detail the history of some early described European Cixiidae, e.g., Reptalus panzeri, R. quinquecostatus, R. melanochaetus and Pentastiridius leporinus and two junior synonyms of P. leporinus (Flata pallens Germar and Flata pallida Herrich-Schäffer, 1835), and make reference to original early works, together with English translations for the first time (see Appendices 1–8). Following the discovery that the type series of R. quinquecostatus is R. melanochaetus of authors and that the British R. panzeri is the same as R. quinquecostatus of continental authors, we provide notes and figures on the types of the former (see Appendix 9) and argue for the retention of the current identity of these two species in continental Europe.

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Resulting from our translation of the introduction of FIEBER’s (1875–79) ‘The Cicadines of Europe’ his unpublished figures from this work were found in the Muséum national d’Histoire naturelle, Paris. These figures include the external male genitalia of Reptalus limbatus (Fieber, 1876) comb. nov., which are found to be very similar to R. venosus (Rambur, 1840) comb. nov. Detailed figures of the former are given here for the first time (see Appendix 10). A revised key to the genera of British Cixiidae and figures of the two British Reptalus species are given (see Appendix 11) together with a brief summary of the European distribution and notes on the ecology of R. quinquecostatus (of authors) (see Appendix 12). Lastly, FIEBER’s (1875) early mention of the word ‘type’ (see the list of his borrowed material, Appendix 7a) prompted us to research the history of the ‘type concept’ and the earliest usage of the word ‘type’. As we found this subject to be poorly documented we give the results of this work in Appendix 13.

Material and methods The following abbreviations are used throughout the text: BMNH ...... The Natural History Museum, London, MNHN ...... Muséum national d’Histoire naturelle, Paris, France ZML ...... Zoological Museum Lemburg, Lvov, Ukraine

Historical account of some early misidentified European Cixiidae Confusion with respect to the identity of some European cixiid species, including the two British Reptalus species, can be traced back to a misidentification of one of the first described European cixiids, Pentastiridius leporinus. This species, described as leporina from , was described after its fluffy (hare-like) wax tail, a characteristic of nymphs and females in this group (LINNAEUS 1761, see Plate 1 and Figs 11a, 11c). A subsequent reference to C. leporina was given by PANZER (1799) who gave a short extract from Linnaeus’s description (Appendix 1) and figured the species for the first time (Fig. 1). His illustration matched the details given by Linnaeus, particularly its fluffy tail and showed a specimen with a short crown, a detail that would be of later significance. Some-time later, HERRICH-SCHÄFFER (1835) described the cixiid Flata pallida (locality unknown) (Appendix 2), and stated in 1837 that his species differed from C. leporina in having a narrower head and in 1838 figured a variety with such a head (HERRICH-SCHÄFFER 1835–1840; Fig. 2). These actions made it possible for SCOTT (1870, in key) to describe the two British cixiid species with five scutellar keels as Cixius leporinus (with ‘crown transverse’) and C. pallidus (with ‘crown longer than broad’). FIEBER (1876) undertook a complete revision of the entire European Auchenorrhyncha fauna (see Appendix 3), during which time he placed and redescribed several cixiids (with five scutellar keels) in Oliarus, including O. leporinus auctt., O.

60 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Identity of some early described European Cixiidae pallidus and O. quinquecostatus (described by DUFOUR 1833 as Cixius quinquecostatus) and also described a new species, Oliarus melanochaetus (see Appendices 4 and 5). The acceptance of PANZER’s (1799) concept of leporinus, by the above authors, was eventually questioned by LÖW (1883) (see Appendix 6), using the works of two Finnish authors. Firstly from SAHLBERG (1871) he learned that Herrich-Schäffer’s description of F. pallida matched that of Linnaeus’s Swedish C. leporina and that the C. leporina of authors (with a short crown) was not to be found anywhere in Scandinavia. Secondly, from REUTER (1880), Löw understood that Herrich-Schäffer’s pallidus is the only species of Oliarus (with five scutellar keels) present in the whole of Scandinavia. He therefore concludes that leporina and pallidus are the same species and that the leporina of other authors requires a new name. For this purpose, Löw chose the name panzeri, after the man who has first figured it, PANZER (1799). In Britain, EDWARDS (1894) followed LÖW’s (1883) work and changed the name of the British pallidus to leporina and leporina to panzeri. This change was subsequently followed by LE QUESNE (1960), who figured their male genitalia. However, although LE QUESNE (1960) used the name panzeri he also noted (p. 8) that it did not match Dlabola’s concept of it (presumably DLABOLA 1952 and 1954 figures) and therefore the name of the British species might have to change.

Results As noted above (see Introduction), the discovery that British Reptalus specimens matched the figures of panzeri in LE QUESNE’s (1960) British handbook, but also matched the figures of Oliarus/Reptalus quinquecostatus given by continental authors, led us to consider the identities of these two species and this led to further discoveries from the pertinent historical literature. The results of these findings are as follows.

Pentastiridius leporinus, P. pallens and P. pallidus With respect to P. leporinus there has been no disagreement as to its identity in recent times (see Fig. 11a). However, only a fragment of the type remains in the Linnaean collection, London, so there is no possibility of confirming its identity. The recent synonymy of Flata pallens Germar, 1821 and Flata pallida Herrich- Schäffer, 1835 with Pentastiridius leporinus, e.g., HOLZINGER et al. (2003), could not be confirmed as the whereabouts of the type material of the latter is unknown and the type of the former is missing its genitalia; in this group the male genitalia are needed for confirmation of species identity (see EMELJANOV 1979). Oliarus pallens (Germar), is treated as a good species in METCALF’s (1936) catalogue with Flata pallida Herrich- Schäffer as its junior synonym, however, if the former is a valid species then it would be a junior homonym of Flata pallida Say, 1830. The original descriptions of F. pallida and F. pallens are provided (Appendices 2 and 7 respectively), together with an image of the type of F. pallens (Fig. 7b). From this and other images of the type of F. pallida seen the five mesonotal keels and large number of spines on the apex of the first hind tarsomere indicate its correct placement in Pentastiridius (see key in HOLZINGER 2003: 70, and

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 61 M. D. WEBB ET AL.

Appendix 11) . It should be noted that METCALF (1936) lists Taurus Mts (Turkey) for the type locality of Flata pallens Germar but the original description says Tauria which is in Crimea, Ukraine.

Reptalus panzeri

As noted above, LÖW (1883), in establishing Oliarus panzeri, was merely renaming what he considered to be the misidentified C. leporina Linnaeus of previous authors and the name he chose was for the ‘discoverer’ of the species, PANZER (1799). Therefore, it could be argued that the original Panzer specimen(s) should be regarded as the type material, but Panzer’s collection has not been found (HORN et al. 1990). Although only three of the five fine mesonotal keels, associated with Reptalus species (including panzeri auctt.), are shown in Panzer’s figure (see Fig. 1), the current identity of panzeri, first figured by DLABOLA (1952, 1954) and more recently by BERTIN et al. (2010), based on male genitalia of non-type material, is upheld.

Reptalus melanochaetus

In describing Oliarus melanochaetus, FIEBER (1876) had only female specimens obtained from Southern France ‘Montpellier, received under the name Fulgora leporina’and Southern Russia ‘Sarepta (Frey-Gessner)’. It was distinguished from its congeners externally by its dark tuberculate setae of the forewing veins and hence the origin of its name (see Appendices 4 and 5). A female specimen from Sarepta has been found in the Paris museum (MNHN, Lethierry-Nouhalier collection) (pers. com. by Gernot Kunz) which could be one of the types. In the absence of evidence to the contrary, the identity of this species given by LOGVINENKO (1975) and subsequent authors is here supported.

Reptalus quinquecostatus With respect to Cixius quinquecostatus Dufour, 1833, there are three specimens standing under this name in Dufour’s collection (MNHN) and as these do not disagree with the original description they are regarded as putative types (see Appendices 8 and 9). Examination of the genitalia of the male specimen for the first time (Figs 9c, d) reveals its identity not to be R. quinquecostatus auctt., as supposed, but identical to R. melanochaetus (Fieber, 1876), sensu LOGVINENKO (1975) and subsequent authors. The other two specimens could be the same species as the syntype male but the tubercle spines of the forewing, which are dark in this species, are rubbed off.

Reptalus limbatus and R. venosus Resulting from our discovery, during the current work, of Fieber’s unpublished figures of his ‘Cicadines of Europe’ (FIEBER 1875–79) the figures of Oliarus limbatus Fieber (Fig. 5d) were found to be similar to those of Cixius venosus Rambur, 1840 given by WEBB (1979: Figs 50–55). This similarity was confirmed by examination of the type

62 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Identity of some early described European Cixiidae specimens of C. venosus (BMNH), with contemporary specimens identified as O. limbatus (MNHN, Puton collection) by Gernot Kunz and Hannelore Hoch who also kindly supplied the figures reproduced here (Appendix 10). The original type data for O. limbatus: ‘Male. 6.5 mm. Body 5 mm. Andalusia, collected by Staudinger (Mus. Vien. n° 166)’ indicated a single specimen in the Natural History Museum, Vienna, but this specimen could not be found there (pers. com. Herbert Zettel). We defer synonymising the two species until further material from other localities is consulted as the slight differences in the configuration of aedeagal spines could be either specific or variation of a single species. However, both species are found to belong in Reptalus based on the five keels on the mesonotum and reduced number of spines on the apex of the first hind tarsomere (see key in HOLZINGER 2003: 70, and Appendix 11), and are thus new combinations in this genus: Reptalus limbatus comb. nov. and R. venosus comb. nov.

Summary and conclusions As the earliest descriptions of many European Cixiidae comprise only details of the external appearance, it is not clear how the identities of some species have been established, as species separation is now reliant mainly on the male genitalia. For example, the original description of Reptalus panzeri could ‘fit’ any cixiid with a short vertex and that of R. quinquecostatus could ‘fit’ any Pentastirini species, i.e. species with five mesonotal keels (EMELJANOV 1971). Even in the monumental work of FIEBER (1876), where an astonishing amount of material was borrowed, including many types, and descriptions of the male genitalia of most cixiids are given in detail, the identities of some species were confused and in fact confusion about the idenity of R. quinquecostatus and R. panzeri, has continued to this day. This includes the identity of one of the two British cixiids with five mesonotal keels, previously known as R. panzeri (= R. quinquecostatus, of authors) (Figs 11d–f). With respect to the identities of R. quinquecostatus and R. melanochaetus, the discoveries outlined in this work require one of two courses of action. Either the type of Cixius quinquecostatus is disregarded and a neotype designated, under the Plenary Powers of the Zoological Commission, Article 75.6 (ICZN 1999; see below), or a new name is given for the species previously identified as R. quinquecostatus in Continental Europe and R. panzeri in Britain. However, with respect to the latter course, as there is no name available a new species would need to be described and also this action would require the synonymy of R. melanochaetus with R. quinquecostatus. The purpose of Article 75.6 of the ICZN (1999) is to promote stability where an identity has been widely accepted for many years. This article states: “When an author discovers that the existing name-bearing type of a nominal species-group taxon is not in taxonomic accord with the prevailing usage of names and stability or universality is threatened thereby, he or she should maintain prevailing usage [Art. 82] and request the Commission to set aside under its plenary power [Art. 81] the existing name-bearing type and designate a neotype. Example. On discovering that the only existing type specimen of Aradus caucasicus Kolenati, 1857 (Heteroptera) was a specimen of another species, KERZHNER & HEISS (1993) proposed that the prevailing usage of the names of both

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 63 M. D. WEBB ET AL. species should be conserved by the designation of a neotype for A. caucasicus under the Commission’s plenary power, and this was accepted in Opinion 1783 (1994)”. In the current case it is the identity of R. quinquecostatus and R. melanochaetus that would be affected if the type of the former is accepted. However, one could also argue that the problem of misidentification of these species has come about through poor taxonomic work, as no one has previously thought to question the identity of R. quinquecostatus and examine the male genitalia of its type, and that such practice should not be condoned. We also feel that disregarding a type should not be undertaken lightly. Therefore has enough time elapsed since the first misidentification and has there been enough published work with the misidentification? Although the ICZN (1999) does not give any advice on this issue we feel sure that on both accounts the currently accepted identities should be upheld. The first misidentification of R. quinquecostatus, was by FIEBER (1876, in key and unpublished figures), followed by the figures of DLABOLA (1952: Figs 11–14), MITYAEV (1971: Plate 7, Figs 13, 14, Plate 8, Fig. 1), LOGVINENKO (1975: Plate 42, Figs 1–3), HOLZINGER et al., (2003: Fig. 58), BIEDERMANN & NIEDRINGHAUS (2004: 120, in pictorial key) and BERTIN et al. (2010: Fig. 3a). Other numerous references to R. melanochaetus and R. quinquecostatus in the literature probably also follow this wrong identity. Based on the above findings we recommend that an application be made to the Zoological Commission to conserve the prevailing usage of the above names, i.e. to disregard the types of Oliarus quinquecostatus, and to erect a neotype (Article 75.6, ICZN 1999). Our discovery that British specimens of Reptalus panzeri are misidentified and are in fact R. quinquecostatus (of continental authors) requires that this name change is made to the British check-list (Michael Wilson & Alan Stewart, in prep.) while here we also provide an update to the key to British Cixiidae (Appendix 11) given by LE QUESNE (1960). Figures of R. panzeri are also provided to help separate it from the other British cixiids with five keels on the mesonotum (P. leporinus). We also give notes on the distribution and ecology of R. quinquecostatus (Appendix 12). The further discovery made, during the current work, of Fieber’s unpublished figures in MNHN that accompanied his (FIEBER 1875–79) ‘The Cicadines of Europe’ sheds light on some other European species that he described. This includes Reptalus limbatus (Fieber, 1876) comb. nov., described from Spain. Fieber’s figures (Fig. 5d) and those supplied by H. Hoch of a male in Puton’s collection (MNHN, Fig. 10) which match a putative type in MNHN (pers. com. Gernot Kunz) show that this species may be the same as Reptalus venosus (Rambur, 1840), comb. nov., also from Spain, the type of which (BMNH, London) was figured by WEBB (1979: Figs 50–55). Subsequent to the finding of Fieber’s manuscript figures those in the family have also been published (GNEZDILOV et al. 2011) and others are being studied by Gernot Kunz (in prep.). Finally, FIEBER’s (1875) early mention of the word ‘Type’ (see his source of material examined, Appendix 3b) prompted us to research the history of the ‘type concept’ and the earliest usage of the word ‘Type’ (see Appendix 13).

64 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Identity of some early described European Cixiidae

Acknowledgements We offer sincere thanks to the many people who have helped to make this study possible. In particular, we wish to thank Hannelore Hoch for revising the manuscript and for making available her drawings of Oliarus limbatus Fieber (see Appendix 10), Mike Wilson and Bill Dollling for their valuable comments, Werner Holzinger for providing information on European Cixiidae and Dufour’s collection, Adeline Soulier-Perkins for the loan of Dufour collection material (MNHN), Roland Mühlethaler and Adeline Soulier-Perkins for (from our enquiries) finding Fieber’s original figures and providing images of Fieber’s plates, Alex Holovachov for type images of Flata pallens Germar and finally Tristan Bantock and Paul Brock for providing their field images of British Cixiidae (see Fig. 11).

References

BERTIN S., PICCIAU L., ACS Z., ALMA A. & BOSCO D. 2010: Molecular differentiation of four Reptalus species (Hemiptera: Cixiidae). Bulletin of Entomological Research 100(5): 551–558. BIEDERMANN R. & NIEDRINGHAUS R. 2004: Die Zikaden Deutschlands. WABV Fründ, Scheeßel, 409 pp. BONAPARTE C. L. 1838: Geographical and comparative list of the birds of Europe and North America, John van Voorst, London, 67 pp. Also available online from: http://www.biodiversitylibrary.org/item/ 103671#page/7/mode/1up. BOURGOIN T. 2013: FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha. Version 8, updated [2013-02-25]. Available online from: http://hemiptera- databases.org/flow/. BUFFON (COMTE DE) G.-L. & DAUBENTON L.-J.-M. 1853: Histoire naturelle générale et particulière: avec la description du Cabinet du Roy. Tome quatrième. Imprimerie royale, Paris, 544 pp. Also available online from: http://gallica.bnf.fr/ark:/12148/bpt6k97493f.r=Buffon+daubenton.langEN BUFFON (COMTE DE) G.-L. 1775: Histoire naturelle des oiseaux. Tome troisième. In: Histoire aturelle, générale et particulière, avec la description du cabinet du roi. Tome dix-huitième. Imprimerie royale, Paris, 502 pp. Also available online from: http://gallica.bnf.fr/ark:/12148/bpt6k975079/f154.image. BUFFON (COMTE DE) G.-L. 1778: Histoire naturelle des oiseaux. Tome quatrième. In: Histoire aturelle, générale et particulière, avec la description du cabinet du roi. Tome dix-neuvième. Imprimerie royale, Paris, 590 pp. Also available online from: http://gallica.bnf.fr/ark:/12148/bpt6k97508n. CZN (COMMITTEE ON ZOOLOGICAL NOMENCLATURE) 1843: Report of a Committee appointed “to consider of the rules by which the Nomenclature of Zoology may be established on a uniform and permanent basis.” In: Report of the Twelfth Meeting of the British Association for the Advancement of Science; held at Manchester in June 1842, 105-121. John Murray, London. Also available online from: http://www.biodiversitylibrary.org/item/46628#page/145/mode/1up. DEMIR E. 2008: The Fulgoromorpha and Cicadomorpha of Turkey. Part I: Mediterranean region (Hemiptera). Munis Entomology & Zoology 3(1): 447–522. Also available online from: http://www.munisentzool.org/ yayin/vol3/issue1/447-522.pdf. DLABOLA J. 1952: Einige neue Paläarktische Zikaden und andere faunistische Bemerkungen. Acta Entomologica Musei Nationalis Pragae 28: 27–37. DLABOLA J. 1954: Køísi – Homoptera. Fauna CSR 1. [Leafhoppers – Homoptera. Fauna of the Czechoslovak Republic, Vol. 1]. Nakladatelství ÈSAV, Praha, 339 pp (in Czech, Russian and German summary). DLABOLA J. 1956: Faunistika a nìkteré nové druhy palearktických køísù. Faunistik und neue Arten der palearktischen Zikaden (Homoptera, Auchenorrhyncha). Acta Entomologica Musei Nationalis Pragae 30 (1955): 121–128 (in German, Czech introduction). DROBNJAKOVIÆ T., PERIÆ P., MARÆIÆ D., PICCIAU L., ALMA A., MITROVIÆ J. & DUBUK B. 2010: Leafhoppers and cixiids in phytoplasma-infected carrot fields: species composition and potential phytoplasma vectors. Pesticidi i Fitomedicina 25: 311–318. Also available online from: http://dx.doi.org/10.2298/PIF1004311D.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 65 M. D. WEBB ET AL.

DUFFIELD C. A. W. 1960: The Auchenorrhyncha (Homoptera) of Kent. Transactions of the Kent Field Club 1(4): 161–170. DUFFIELD C. A. W. 1963: The genus Aphrodes (Homoptera: Auchenorrhyncha). Transactions of the Kent Field Club 1(4): 155–160. DUFOUR L. 1833: Recherches anatomiques et physiologiques sur les Hémiptères, accompagnées de considérations relatives à l’histoire naturelle et à la classification de ces insectes. Mémoires des Savants étrangers de l’Académie Royale des Sciences de l’Institut de France 4: 129–462, plates: 1–19. DUFOUR L. 1839: Quelques observations sur une note de M. Doyère relative au tube digestif des cigales. Annales des Sciences Naturelles, Zoologie et Paléontologie comprenant l’Anatomie, la Physiologie, la Classification et l’Histoire Naturelle des Animaux 12(2): 287–289. Also available online from: http://www.biodiversitylibrary.org/item/47937#page/293/mode/1up. EDWARDS J. 1894: The Hemiptera Homoptera of the British Islands. A descriptive account of the families, genera, and species indigenous to Great Britain and Ireland, with notes as to localities, habitats, etc. Vol. 1. L. Reeve & Co., London, 32 pp. EMELJANOV A. F. 1971: New genera of planthoppers of the fauna of the USSR from families Cixiidae and Issidae (Homoptera, Auchenorrhyncha). Entomological Review 50(3): 619–627. EMELJANOV A. F. 1978: (New genera and species of leafhoppers (Homoptera, Auchenorrhyncha) from the USSR and Mongolia). Entomologicheskoe Obozrenie 57(2): 316–332 (in Russian, English translation published in Entomological Review 57(2): 220–233). FARBER P. L. 1976: The Type-Concept in Zoology during the First Half of the Nineteenth Century. Journal of the History of Biology 9(1): 93–119. http://www.jstor.org/stable/4330645. FENALD H. T. 1939: On Type Nomenclature. Annals of the Entomological Society of America 32: 689. FIEBER F. X. 1875: Les Cicadines d’Europe d’après les originaux et les publications les plus récentes. Première partie. Revue et Magasin de Zoologie 3(3): 288–416. Also available online from: http://www. biodiversitylibrary.org/item/82428#page/9/mode/1up. FIEBER F. X. 1876: Les Cicadines d’Europe d’après les originaux et les publications les plus récentes. Deuxième partie. Revue et Magasin de Zoologie 4(3): 11–268. Also available online from: http://www. biodiversitylibrary.org/item/82428#page/139/mode/1up. FIEBER F. X. 1878: Les Cicadines d’Europe d’après les originaux et les publications les plus récentes. Deuxième partie (suite). Revue et Magasin de Zoologie 5(3): 1–45. Also available online from: http://www. biodiversitylibrary.org/item/82428#page/397/mode/1up. FIEBER F. X. 1879: Les Cicadines d’Europe d’après les originaux et les publications les plus récentes. Troisième partie. Revue et Magasin de Zoologie 6(3): 270–308. Also available online from: http://www. biodiversitylibrary.org/item/82428#page/437/mode/1up. FRIZZEL D. 1933: Terminology of Types. The American Midland Naturalist 14(6): 637–668. GERMAR E. F. 1821: Bemerkungen über einige Gattungen der Cicadarien. Magazin der Entomologie 4: 1–106. Also available online from: http://www.biodiversitylibrary.org/item/81944#page/5/mode/1up. GJONOV I. 2004: Cicadina (Insecta: Homoptera) of the Eastern Rhodopes (). Pp. 315–330. In: BERON P. & P OPOV A. (eds.): Biodiversity of Eastern Rhodopes (Bulgaria and Greece). Pensoft Publishers, Sofia, 951 pp. GNEZDILOV V. M., SOULIER-PERKINS A. & BOURGOIN T. 2011: Fieber’s original drawings and their corresponding types for the family Issidae (Hemiptera, Fulgoromorpha) in the Museum national d’Histoire naturelle of Paris, France. Zootaxa 2806: 24–34. GRAY G. R. 1840: A list of the genera of birds, with an indication of the typical species of each genus compiled from various sources. [London]: R. and J. E. Taylor. viii+ii+80 pp. Also available online from: http://dx.doi.org/10.5962/bhl.title.13777. HERRICH-SCHÄFFER G. A. W. 1835–1840: Homoptera. In: Nomenclator entomologicus: Verzeichniss der europäischen Insecten, zur Erleichterung des Tauschverkehrs mit Preisen versehen. Pustet, Regensburg, 116 + 244 pp. Also available online from: http://www.biodiversitylibrary.org/item/78852. HOLZINGER W. E. 2009: Auchenorrhyncha (Insecta). Pp. 41–100. In: SCHUSTER R. (ed.): Checklisten der Fauna Österreichs, No. 4. Biosystematics and Ecology Series No. 26, Austrian Academy of Sciences Press, Vienna, 100 pp. HOLZINGER W. E., EMELJANOV A. F. & KAMMERLANDER I. 2002: The family Cixiidae Spinola 1839 (Hemiptera: Fulgoromorpha) – a review. Pp. 113–138. In: HOLZINGER W. E. (ed.): Zikaden leafhoppers, planthoppers and cicadas (Insecta: Hemiptera: Auchenorrhyncha). Denisia 4: 1–556.

66 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Identity of some early described European Cixiidae

HOLZINGER W. E., KAMMERLANDER I. & NICKEL H. 2003: Fulgoromorpha, Cicadomorpha excl. Cicadellidae: The Auchenorrhyncha of Central Europe/Die Zikaden Mittleuropas. E. J Brill, Leiden, 673 pp. HORN W., KAHLE I., FRIESE G. & GAEDIKE R. 1990: Collectiones Entomologicae: Ein Kompendium über den Verbleib entomologischer Sammlungen der Welt bis 1960. Akademie der Landwirtschaftswissenschaften der Deutschen Demokratischen Republik, Berlin, 2 parts, 573 pp. ICZN (INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE) 1941: Interpretation of the expression “typical species” in Westwood’s (1840) Synopsis. Smithsonian Miscellaneous Collections 73: 16–18. Also available online from: http://www.biodiversitylibrary.org/item/35815#page/30/mode/1up. ICZN (INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE) 1999: International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature c/o the Natural History Museum, London, 271 pp. Also available online from: http://iczn.org/iczn/index.jsp. ICZN (INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE) 1994. Opinion 1783. Aradus caucasicus Kolenati, 1857 (Insecta, Heteroptera): syntype replaced by a neotype, so conserving the usage of the specidfic name and that of A. hieroglyphicus Sahlberg, 1878. Bulletin of Zoological Nomenclature 51(3): 279. Also available online from: https://archive.org/stream/bulletinofzoolog51inte#page/278/mode/2up. JONES R. A. & HODGE P. J. 1999: Notes on the occurrence of the planthoppers Reptalus panzeri (Löw) (Hemiptera: Cixidae) and Asiraca clavicornis (Fab.) (Hemiptera: Delphacidae). British Journal of Entomology and Natural History 12: 239–240. Also available online from: http://www.biodiversitylibrary. org/item/94410#page/281/mode/1up. KERZHNER I. M. & HEISS E. 1993: Aradus caucasicus Kolenati, 1857 (Insecta, Heteroptera): proposed replacement of syntype by a neotype, so conserving usage of the specific name and that of A. hieroglyphicus Sahlberg, 1878. Bulletin of Zoological Nomenclature 50(2): 115–117, Case 2843. Also available online from: http://www.biodiversitylibrary.org/page/12249540#page/137/mode/1up. KIRBY W. & SPENCE W. 1826: An Introduction to entomology: or elements of the natural history of insects: with plates, vol. 4. Longman, Rees, Orme, Brown, and Green, London, 634 pp. Also available online from: http://www.biodiversitylibrary.org/item/79774#page/11/mode/1up. KIRBY P. 1992: A review of the scarce and threatened Hemiptera of Great Britain. UK Nature Conservation: 2. Joint Nature Conservation Committee, Peterborough, 267 pp. KIRBY P. 2001: Habitat Management for Invertebrates. A practocal Handbook. Royal Society for the Protection of Birds, Sandy, 150 pp. KUSNEZOV V. 1937: Neue asiatische Oliarus-Arten (Homoptera Cixiidae). Entomologisches Nachrichtenblatt 10: 161–168. LAMARCK J. B. P. A. DE M. 1801: Système des animaux sans vertèbres ou tableau général des classes, des ordres et des genres de ces animaux; présentant leurs caractères essentiels et leur distribution, d’après la considération de leurs rapports naturels et de leur organisation, et suivant l’arrangement établi dans les galleries du Muséum d’Hist. Naturelle, parmi leurs dépouilles conservées; précédé du discours d’ouverture du cours de Zoologie donné dans le Muséum National d’Histoire Naturelle l’an 8 de la Republique. ‘An IX’. Auteur & Déterville, Paris, VIII+432 pp. Also available online from: http://www.lamarck.cnrs.fr/ice/ice_page_detail.php?lang=fr&type=text&bdd=lamarck&table=ouvrages_l amarck&bookId=7&typeofbookDes=Livres&pageOrder=4&facsimile=off&search=no. LATREILLE P. A. 1802: Histoire naturelle, générale et particulière des Crustacés et des Insectes: ouvrage faisant suite aux oeuvres de Leclerc de Buffon, et partie du cours complet d’histoire naturelle rédigé par C. S. Sonnini, membre de plusieurs Sociétés savantes. Vol. 3: i–xii, F. Dufart, Paris, pp. 13–467. Also available online from: http://www.biodiversitylibrary.org/item/80064#page/5/mode/1up. LATREILLE P. A. 1804: Histoire naturelle, générale et particulière des Crustacés et des Insectes: ouvrage faisant suite aux oeuvres de Leclerc de Buffon, et partie du cours complet d’histoire naturelle rédigé par C. S. Sonnini, membre de plusieurs Sociétés savantes. Vol.7. F. Dufart, Paris, pp. 5–413, plates lviii–lxvi. Also available online from: http://www.biodiversitylibrary.org/item/80060#page/5/mode/1up. LAUTERER P. 1957: Pøíspìvek k poznání køísù ÈSR (Hom., Auchenorrhyncha). [Contribution à la connaissance des Cicadines de la Tchécoslovaquie (Hom., Auchenorrhyncha).] Acta Societatis Entomologicae Èechosloveniae 54: 1–3 (in Czech, French summary). LE QUESNE W. J. 1960: Hemiptera-Homoptera: Fulgoromorpha. Handbooks for the Identification of British Insects. Volume 2 (3). Royal Entomological Society, London, 68 pp.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 67 M. D. WEBB ET AL.

LE QUESNE W. J. & PAYNE K. R. 1981: Cicadellidae () with a checklist of the British Auchenorrhyncha. Handbooks for the Identification of British Insects. Volume 2 (2c). Royal Entomological Society, London, 195 pp. LINNAEUS C. 1761: II. Hemiptera. Cicada. In: Fauna svecica: sistens animalia sveciae regni: Mammalia, Aves, Amphibia, Pisces, Insecta, Vermes, distributa per classes & ordines, genera & species, cum differentiis specierum, synonymis auctorum, Editio altera, auctior. insectorum. L. Salvii, Stockholmiae, 578 pp., plates 1–2. Also available online from: http://www.biodiversitylibrary.org/item/100333. LINNAVUORI R. E. 1965: Studies on the South- and East-Mediterranean Hemipterous fauna. Acta Entomologica Fennica 21: 1–69. LÖW P. 1883: Zur synonymie der Cicadinen. Wiener Entomologische Zeitung 2: 147–150. Also available online from: http://www.landesmuseum.at/pdf_frei_remote/WEZ_2_0147-0150.pdf. LOGVINENKO V. N. 1975: Fulgoroidny cikadovy Fulgoroidea. [Planthoppers Fulgoroidea.] Fauna Ukrainy, Kyiv 20(2): 1–287 (in Ukrainian). MALENOVSKÝ I., BAÒAØ P. & KMENT P. 2011: A contribution to the faunistics of the Hemiptera (Cicadomorpha, Fulgoromorpha, Heteroptera, and Psylloidea) associated with dry grassland sites in southern Moravia (Czech Republic). Acta Musei Moraviae, Scientiae biologicae (Brno) 96(1): 41–187. MALENOVSKÝ I. & LAUTERER P. 2012: Leafhoppers and planthoppers (Hemiptera: Auchenorrhyncha) of the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). In: MALENOVSKÝ I., KMENT P. & KONVIÈKA O. (eds.): Species inventories of selected insect groups in the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). Acta Musei Moraviae, Scientiae biologicae (Brno) 96(2) (2011): 155–322. MCCOY F. 1876: Decade III. Prodromus of the palaeontology of Victoria; or Figures and descriptions of Victorian Organic remains. Skinner, Melbourne & Trübner, London, 40 pp. Also available online from: http://www.biodiversitylibrary.org/item/51044#page/11/mode/1up. MELICHAR L. 1896: Cicadinen (Hemiptera-Homoptera) von Mittel-Europa. Felix L. Dames, Berlin, 364 pp. Also available online from: http://www.biodiversitylibrary.org/item/35346. METCALF Z. P. 1936: Fascicle IV Fulgoroidea. Part 2. Cixiidae. In: General catalogue of the Hemiptera. Vol. 4(2). Smith College, Northhampton, MA, 269 pp. Also available online from: http://www.biodiversitylibrary.org/item/29871#page/3/mode/1up. MITYAEV I. D. 1971: Cicadovye Kazakhstana (Homoptera-Cicadinea). [Leafhoppers of Kazakhstan (Homoptera-Cicadinea).] Nauka, Alma-Ata, 210 pp (in Russian). MOZAFFARIAN F. & WILSON M. R. 2011: An annotated checklist of the planthoppers of Iran (Hemiptera, Auchenorrhyncha, Fulgoromorpha) with distribution data. ZooKeys 145: 1–57. Also available online from: http://dx.doi.org/10.3897/zookeys.145.1846. PANZER G. W. F. 1799: Fulgora virescens, Fulgora obliqua, Cicada costata, Cicada cruentata, Cicada haemorrhoa, Cicada aethiops, Cicada thoracica, Cicada leporina. Faunae Insectorum Germanicae initia: oder Deutschlands Insecten 61: 12–19. Also available online from: http://www.biodiversitylibrary.org/ item/53215#page/79/mode/1up. PINZAUTI F., TRIVELLONE V. & BAGNOLI B. 2008: Ability of Reptalus quinquecostatus (Hemiptera: Cixiidae) to inoculate stolbur phytoplasma to artificial feeding medium. Annals of Applied Biology 153: 299–305. PRUTHI H. S. 1930: Studies on Indian Jassidae (Homoptera). Part 1. Introductory and description of some new genera and species. Memoirs of the Indian Museum 11: 1–68. RAMBUR J .P. 1840: Seconde section des Hémiptères. Les Homoptères Latreille. Faune entomologique de l’Andalousie 5: 177–304. RAMSAY A. 2009: Invertebrate Survey, Stoke to Yantlet Pit, Isle of Grain, Kent. In: Stoke to Yanlet Pit Report to The Oil and Pipelines Agency. RSK Carter Ecological, Banbury, pp. 39–44. RÉAUMUR R. F. DE 1747: Différens Moyens d’empêcher de se corrompre les Oiseaux morts qu’on veut envoyer dans des Pays éloignez, & de les y faire arriver bien conditionnez. Quelques-uns de ces mêmes moyens peuvent être aussi employez pour conserver des Quadrupèdes, des Reptiles, des Poissons et des Insectes. S. 1: [Imprimerie Royale] (1747), in-4, 4 pp. Also available online from: http://www.pierre- poivre.fr/reaumur-2-prospectus.pdf. RÉAUMUR R. F. DE 1748: Divers means for preserving from corruption dead birds, intended to be sent to remote countries, so that they may arrive there in good condition. Some of the same means may be employed for preserving quadrupeds, reptiles, fishes, and insects. Translated from the French by Phil. Hen. Zollman, Esq;

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F.R.S. Philosophical Transactions of the Royal Society of London 45: 305. Also available online from: http://dx.doi.org/10.1098/rstl.1748.0034. REUTER O. M. 1880: Nya bidrag till Åbo och Ålands skärgårds Hemipter-fauna. Meddelanden af Societas pro Fauna et Flora Fennica 5: 160–236. Also available online from: http://www.biodiversitylibrary. org/item/23882#page/168/mode/1up. SAHLBERG J. R. 1871: Öfversigt af Finlands och den Skandinaviska hälfons Cicadariae. I. Notiser ur Sällskapets pro Fauna et Flora Fennica Förhandlingar (N.S.) 9(12): 1–506, plates 1–2. Also available online from: http://www.biodiversitylibrary.org/item/47587#page/7/mode/1up. SAY T. 1830: Descriptions of new North American hemipterous insects, belonging to the first family of the section Homoptera of Latreille. Journal of the Academy of Natural Sciences of Philadelphia 6: 235–244. Also available online from: http://www.biodiversitylibrary.org/item/79409#page/247/mode/1up. SCHUCHERT C. 1897: What is a type in natural history? Science 5(121): 636–640. Also available online from: http://www.jstor.org/stable/1624022. SCOTT J. 1870: On certain British Hemiptera-Homoptera. Revision of the family Cixiidae. Entomologist’s Monthly Magazine 7: 118–123. Also available online from: http://www.biodiversitylibrary.org/ item/102841#page/152/mode/1up. SELJAK G. 2004: Contribution to the knowledge of planthoppers and leafhoppers of Slovenia (Hemiptera: Auchenorrhyncha). Acta Entomologica Slovenica 12(2): 189–216. STRICKLAND H. E. 1837: Rules for Zoological Nomenclature. The Magazine of Natural History and Journal of Zoology, Botany, Mineralogy, Geology, and Meteorology (N. S.) 1: 173–176. Also available online from: http://www.biodiversitylibrary.org/item/87352#page/189/mode/1up. STRICKLAND H. E. 1845: Report on the Recent Progress and Present State of Ornithology. Pp. 170–221. In: Report of the Fourteenth Meeting of the British Association for the Advancement of Science; held at York in September 1844. John Murray, London. Also available online from: http://archive.org/ stream/report29sciegoog#page/n215/mode/2up. STRICKLAND H. E. 1878: Rules for Zoological Nomenclature. John Murray, London. 26pp. Also available from: http://www.archive.org/stream/rulesforzoologi00sciegoog#page/n3/mode/1up. THOMAS O. 1893: Suggestions for the more definitive use of the word “type”. Proceedings of the Zoological Society of London 1893(2): 241–242. Also available online from: http://www.biodiversitylibrary.org/ item/97156#page/315/mode/1up. VAN DER HAMMEN L. 1981: Type concept, higher classification and evolution. Acta Biotheoretica 30(1): 1–48. WALKER F. 1850: List of the specimens of homopterous insects in the collection of the British Museum. London: British Museum. 1: 1–260. Also available online from: http://www.biodiversitylibrary.org/item/ 132966#page/3/mode/1up. WALSINGHAM T. & DURRANT J. H. 1896: Rules for regulating nomenclature: with a view to secure a strict application of the law of priority in entomological work. Longmans, London & New York, 18 pp. WEBB M. D. 1979: Revision of Rambur’s Homoptera species from the types in the British Museum. Annales de la Societé Entomologique de France (N.S.) 15(1): 227–240. WESTWOOD J. O. 1840: Synopsis of the genera of British insects. In: An introduction to the modern classification of insects. Longman, Orme, Brown, Green, Longmans. London. 2: 1–158. Also available online from: http://www.biodiversitylibrary.org/item/45812#page/607/mode/1up. WHEWELL W. 1840: The Philosophy of the Inductive Sciences, founded upon their history. (2 vols), John W. Parker, London, 708 pp. 2nd edition 1847, Also available online from: http://books.google.com/ books?id=um85AAAAcAAJ. WHITE A. 1845: XIV. – Description of an apparently new species of Longicorn from Mexico in the collection of the British Museum. Annals and Magazine of Natural History 15: 108–111. Also available online from: http://www.biodiversitylibrary.org/item/19398#page/134/mode/1up WILLIAMS D. M. & EBACH M. C. 2008: Foundations of systematics and biogeography. Springer, 309 pp. WILSON S. W., MITTER C., DENNO R. F. & WILSON M. R. 1994: Evolutionary Patterns of host plant use by Delphacid Planthoppers and their relatives. Pp. 7–113. In: DENNO R. F. & PERFECT T. J. (eds.): Planthoppers: their ecology and management. Chapman & Hall, London, 800 pp.

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Appendix 1. The description of Cicada leporina Linnaeus, sensu Panzer, 1799 (= Oliarus panzeri Löw, 1883) (original and translation, from PANZER 1799) Note. The following description of PANZER (1799) appeared under LINNAEUS’s (1761) species name “leporina”, meaning hare-like and presumably referring to the fluffy wax ‘tail’. This feature was mentioned in Linnaeus’s description as “woolly anus” and in Panzer’s description as “Federbusch” (a term normally referring to a head plume). These terms together with Panzer’s figure of the female indicate that the description was based on the female. The passage consists of Linnaeus’s original summary to his species description, its original and subsequent references (not cited here), habitat and collector information, and finally a brief comment by Panzer and legend to his figure. The species was subsequently renamed as a new species by LÖW (1883) (see Appendix 6).

Fig. 1. The original description and figure of Oliarus panzeri, from PANZER (1799: 19).

Translation from PANZER (1799) [added text is in square brackets] CICADA leporina. Die Cicade mit dem Federbusch. Cicada leporina: wings bent downwards transparent: in front anastomoses pale above line blackish, anus woolly. ... Habitat in Austria. Dn. de Megerle. In the illustrated Fabricius [1799] this Cicada is not yet made known and everybody everywhere refrains from mentioning it. In my opinion it should be counted among the Cercopids. [Figure legends:] a Actual size. b Enlarged. c Antenna enlarged.

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Appendix 2. The description of Flata pallida Herrich-Schäffer, 1835 (original and translation and his 1838 figure of un-named variety of pallida)

Fig. 2. The original description of Flata pallida (from HERRICH-SCHÄFFER 1835: 64); the original figure of Flata pallida un-named variety (from HERRICH-SCHÄFFER 1838: plate no. 154.4).

Translation of the original description of Flata pallida from HERRICH-SCHÄFFER (1835: 64) 55 Flata. I. Costal veins not spotted 1. Posterior tibiae smooth. musiva 2. - - with spines A. Elytra whitish-hyaline, veins dusky, black-spotted, stigma black- dusky, broadly white at base, first radial cell entering[?!]; apical cells 4, 6 and 9 petiolate. leporinus B. Elytra yellowish-hyaline, veins yellow, dusky-banded toward the apex, stigma white delimited internally by a dusky line; radial cells 2–4, 6 and 9 petiolate. pallidus II. Costal veins spotted [leads to simplex, albicincta, contaminata, stigmatica and nervosa]

Note. Herrich-Schäffer (or the printer) seems to have made a mistake with the last line, omitting the contrast to the radial cell and not stating that it is the apical cells 2–4, 6 and 9 that are petiolate.

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Appendix 3a. Preface of the translator of FIEBER’S (1875) ‘The Cicadines of Europe’ Part 1 (original and translation)

Note. Fieber's 'The Cicadines of Europe' was published posthumously in several parts in French from the original unpublished German manuscript, the later work now located in MNHN. Here we give the Preface by its translator (Reiber) followed by our translation of the same. This first part of Fieber’s study deals with morphology and gives a key to genera. Species descriptions, in the form of a key, formed the basis of the remaining parts of which the Cixiidae were dealt with in the second part (FIEBER 1876). Fieber’s reference to his unpublished figures (found in MNHN as a result of our study), are noted in the third paragraph (see also Appendix 3b).

Fig. 3. Preface of the translator of FIEBER’s (1875) ‘The Cicadines of Europe’ Part 1, pages 288–290.

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Translation of the Preface of the translator of Fieber’s (1875) ‘The Cicadines of Europe’ Part 1, pages 288–290

THE CICADINES OF EUROPE FROM THE ORIGINALS AND THE MOST RECENT PUBLICATIONS FIRST PART: Comprising the families of the Membracida, Cicadaea, Fulgorida, Cercopida, Ulopida, Paropida, Scarida. arranged according to the analytic method. By Dr Franz-Xavier FIEBER ______Translated from German By Ferd. REIBER, Member of the entomological Society of France ______

PREFACE OF THE TRANSLATOR

In 1872, Dr Fieber published the catalogue of the Cicadines of Europe1. This catalogue was the precursor of a big descriptive work that the Austrian scientist intended to publish soon after. Illness first delayed the publication of the announced book, and, finally, on February 23, 1872, the author’s death came to postpone its appearance indefinitely. The editor returned the manuscript to Mrs Fieber, and there was not one German naturalist to be found who wanted to honour the memory of the great hemipterist by taking care of the impression of his work. My friend Dr Puton, alone, constantly made the biggest efforts to save from oblivion this masterpiece of patience. His efforts unfortunately remained fruitless. At the beginning of this year, I decided in my turn to start again the steps where Dr Puton had left them. Luckier than he, I had the satisfaction to save the manuscript. I bought it from the author’s widow sharing the expenses with Messrs Puton, de Remiremont, and Lethierry, of Lille, since a long while in possession of Fieber’s drawings. Unfortunately, some parts of the manuscript were already lost. Placed with the alternative to publish a piecemeal work or to complete it, my friends didn’t hesitate. They offered me to complete it, which meant for them to undertake the monograph of the lost genera while they asked me to translate into French the saved part of the manuscript, in order to be able to publish a homogeneous work in the same language. I first hesitated to undertake this difficult and long-winded task but my friends having offered to me their devoted support, my scruples departed and I set to work. Such is, to sum it up, the history of the work of which we publish the beginning today. We will reproduce the author’s manuscript faithfully, without any innovation and without adding the new described species since the appearance of the catalogue that summarizes the whole work. Only the lost genera, treated again by Messrs Puton and Lethierry, will be an exception to the rule and will be special monographs of their authors. Our goal is to save from oblivion the fruit of our venerated master’s long years of study and to deliver to the entomological public a general work on an order of insects still very little studied due to the lack of general works. May we bring this enterprise to a successful completion! As for me, I am pleased to endow French science with the translation of an unpublished masterly work. I dare hope that French entomologists will forgive me the imperfections of this translation while taking into account the difficulties that present such an enterprise. Ferd. REIBER. Strasbourg, June 1875.

1 Katalog d. europaeischen Cicadinen. Gerold's Sohn, 1872.

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Appendix 3b. Preface by Fieber to his ‘The Cicadines of Europe’ Part 1, 1875 (translation from FIEBER 1875: 290–294) Note. Fieber states below that he was able to borrow specimens (including ‘types’) from all those that he requested material from (except Kirschbaum). He also notes that he figured in colour all the type specimens (see fourth paragraph from end of Preface). Although these figures were not reproduced it would appear from the translator’s comments above '(Appendix 3a) and translators footnote (see end of this Appendix) that they were in the possession of Lethierry, before being given to the Paris Museum (see Appendix 5).

PREFACE OF THE AUTHOR Different friends having asked me to get back to my works on the Cicadines, works interrupted for many years due to the lack of books and sufficient material, such as original specimens, I decided, in the year 1864, to put myself back to the task and to undertake a work on the Cicadines of Europe. I didn’t hide the importance of this enterprise to myself, but I had however no idea of the pain that such a work required or of the difficulties it presented. Indeed, it is only in the course of these new studies that I recognized how difficult it is to gather the necessary materials, often scattered in works and collections not so easily available, to examine and to describe all species and the originals according to a method appropriate to the present level of science, and to guide oneself, in the absence of all other general work, apart from the Rhynchoten Lievlands, by Dr G. Flor, 1861.

[The next section of Fieber’s preface deals with the source of his borrowed material. These we have rearranged alphabetically and paraphrased]. Mr Berquier, indeterminate Cicadines, collected around Trieste [Italy]. Mr Bohemann loaned, when in Prague, a lot of Swedish Cicadines and his Nova Svenska Homopts. Mr Brischke, undetermined Hemiptera and Cicadines, gathered around Danzig [Poland] and exceptionally prepared. Mr Erber, several times, consignments from Corfu, Syra, Tixos, Montenegro, in which were many rarities or novelties Dr Flor sent his new species as well as a lot of others, and his work on the Rhynchotes of Livonia. Knight von Frauenfeld, a lot of small indeterminate Cicadines, for the most part Jassids, and of Austrian source. Mr. Frey-Gessner placed at my disposal his rich collection of species from Switzerland, the south of Russia, Corsica, France and Spain (collected by Meyer-Dür). Mr Fritsch, vice-director, Cicadines from Bohemia and Salzburg Pater Vincent Gredler, Tyrolean species [Austria]. Dr Heller, unnamed species, collected around Innsbrück and in the Stubaithal [Austria].

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Mr Lucas von Heyden, for the communication of all Cicadines that served to the work of Kirschbaum, entitled the “ Cicadinen von Wiesbaden, “ communication which allows me to review and to rectify this author’s work. Messrs Lethierry and Dr Puton, bugs from the department of the North and from Algeria Dr Mayr, a great deal of Herrich-Schaeffer’s types, of Austrian and Hungarian species, of others collected around Naples by Mr Emery, and, finally, Cicadines described by Kirschbaum. Mr Mink communicated me his Cicadines collected around Crefeld [Germany]. Messrs Mulsant and Rey loaned a great deal of Cicadines from the South of France, among which were several of the species described by them, as well as some new species. Dr Nowicky, Hemiptera and Cicadines of the Carpatheans and different regions of the Galicia. Mr Oschannin, Russian species from some provinces. Dr Redtenbacher, director of the Imperial Cabinet of Natural History in Vienna, and of Mr Rogenhofer, its curator, the communication of the typical species of Fabricius, Megerle, Mann and Kolenaty which can be found at this museum. Mr Scott (p. 292) species from different parts of England. Dr Stål was promising to come to my help by procuring for me the types of the Swedish authors, a promise kept by the consignment of these types and of a lot of species from other countries. Dr Stein sent me a great deal of small species collected in Greece by Dr Krüper, species of which some were named; he added to it a certain number of Germar’s types, on the validity of which I had some doubts as species. Messrs Ungerer and Professor Kissl, Hemiptera and Cicadines from Bavaria. Mr Wüstnei, the candidate in philosophy, species from Mecklenburg [Germany].

I extend here to all aforementioned correspondents my best thanks for the confidence they have shown me in sending their Cicadines and in thus allowing me to write and complete the present work. The unfortunate postponement of the appearance of this book comes in part from my serious and long illness of the years 1868 and 1869–1870. While examining the rich materials at my disposition, I recognized that the venation of the elytra and wings was of prime importance in order to separate and create genera, but that the characters taken from the shape of the head, the antennae, from the pro- and mesonotum, the legs and their different armature, also had to be taken into consideration, naturally. As for distinguishing the species, I had to use not only on the characters taken from the structure of the different parts of the body, the outline, the coloration and the armature, but especially the examination of the genitalia of the two sexes. Dr Flor had already recognized partially, in his excellent work on the Rhynchotes of Livonia, that the examination of the genitalia was absolutely necessary for the distinction of the species. Kirschbaum, in this matter, has only imitated him. I found, in addition to the organs mentioned by Dr Flor, / / p. 293 / / two other organs used neither by him, nor by Kirschbaum: 1. the bent, non articulate styles, styli, griffel

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[mentioned by mistake in the Grundzüges zur gener. Theilung d. Delphacini, in the Verhanld.k.k.zool. bot. Ges., 1866, under the name of Raife (hoops, in the Orthoptera, Cerci, articulate anal appendices, griffel)], that one may find among the males, in the open (in the delphacids, cixiids, etc.) or hidden (in the scarids and the jassids). Flor reports these appendices in the Cixius, but is not concerned further about them. Kirschbaum only describes them for some genera of Fulgorids, and often in an inexact way; he does not bother about them with the Cercopids, Jassids. 2. Among the females of the Delphacines, the lateral plates (nebenplatten), paraplagae. [ibid, p. 517; basilary or lateral plates (grundplatten or nebenplatten). J. R. Sahlberg in his Ofvers. Finlands Cicadaria, 1871, t. I, fig. 6, calls them lateral lobi.] These two organs provide, in their different variations, good characters for the exact determination of the species. After grouping and inspection of the materials at my disposition, I found that the number of the species had nearly doubled, more than doubled even in a lot of genera. Once it was begun, I didn’t spare any pains or money to bring the work to a satisfactory conclusion. As – according to Dr Stal – the types of Fallen no longer exists today other than by tradition, as many other species are very difficult to see or to obtain, since unique examples are scattered in various collections and a lot of types are lost due to accident or decay, I reproduced by coloured drawings each of the original types that I had been given to examine. These drawings1, / / p. 294 / / 10 of which appear on every 8°, should serve to distinguish in times to come the species in an indubitable way and to form the basis on which rests the present book. The genera and their characters are featured in a separate notebook, and to the number of 6 on every 8°. The present work will therefore consist of 3 volumes with the drawings. I have a thorough knowledge of the literature dealing with my topic. I am in part indebted for it to Messrs Stål, Mayr, Signoret, Scott, J., Sahlberg and Rogenhofer, for sending excerpts of works in various languages, difficult to obtain, and in loans of the Viennese libraries of the Imperial Cabinet of Zoology, the Court and Society zoolog. Botany. I owe to M. Dohrn the Fulgorids of Dr Schaum (in Encycl. Der Wissensch). Mr. Kirschbaum whom I have, since 1866, asked five times by letter to communicate to me the new species that he personally possesses and which I didn’t see in other collections, has kept an obstinate silence, although I have offered him twice the guarantee for the safe return of his consignment. There remains therefore only a small number of his novelties which I haven’t seen and of that the value as species is doubtful, given that no one knows them. Mr. Kirschbaum probably fears changes to his determinations and finds it more important to publish many new species, without sufficient knowledge of those described hitherto, quickly and without complete knowledge of the bibliography dealing with the topic. This naturalist prefers to leave others to solve his enigmas. The limits of the fauna of the Cicadines of Europe, and the zones where they abound are the same than those of my Hemiptera of Europe. Dr F.-X. FIEBER 1 These coloured drawings --the finish and accuracy of which is admirable-- are now owned by Messrs. Lethierry and Puton who would be happy to make them available to the publisher or the Society willing to publish them.

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Appendix 4. The description of Oliarus melanochaetus Fieber, 1876 (original and translation) Note. Fieber’s description of O. melanochaetus appeared in the first half of the first couplet to a key of Oliarus species.

Fig. 4. The original description (in key) of Oliarus melanochaetus (from FIEBER 1876: 187–188).

Translation of the original description of Oliarus melanochaetus from FIEBER (1876: 187–188). The alternative halves of the couplet are emphasized in bold. 1. Bristles of the granules of the nerves black; granules brown, nerves yellowish. Elytra hyaline. Angular nerves brown with yellow brownish shades; small spots of a yellowish-brown on the forks of the clavus, the interior sector and the apical nerves. Tegula yellow, with a black base. Median carina of the face yellow until above the clypeus; frons black with its linear edge yellow; edge of the clypeus finely yellow. Vertex longer than broad at the front between the eyes. Pronotum yellow, of a black brown under the eyes. Mesonotum black, with the posterior edge and the tip of a yellowish white. Stigma dull, yellowish white, sometimes internally brownish, with some brown granules. Wings hyaline; nerves of their apical half brownish. Abdomen completely black with yellow orange edges. Coxae and femurs black; their extremities, tibiae and posterior tarsi of a yellowish white. Anterior and intermediary tarsi, base of the tibiae and last segment of the posterior tarsi brownish. ♀ 8 mm. South of France, Montpellier, known as Fulgora leporina and southern Russia, Sarepta (Frey-Gessner)...... 1. O. melanochaetus Fieb. – Bristles of the granules yellowish...... 2

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Appendix 5. The unpublished figures that accompanied Fieber’s (1875–1879) ‘The Cicadines of Europe’

Notes. The unpublished figures of FIEBER’s ‘The Cicadines of Europe’ (1875–1879) were found in the Paris Museum Library as a result of the current work. Figures from those plates, showing Oliarus quinquecostatus, O. melanochaetus and O. limbatus, are reproduced here (Fig. 5). The genitalia figures of Fieber’s Fig. 16 are labelled “artemisae” but crossed through and quinquecostatus written in pencil. According to METCALF’s (1936) catalogue Oliarus artemisae Becker is a nomen nudum and synonym of O. quinquecostatus. The only place “artemisae” appears in Fieber’s key is under O. quinquecostatus, saying the two species are confused. The following discrepancies are found with respect to the figure numbers given in FIEBER’s (1876) Cixiidae key and the unpublished figures: O. quinquecostatus is numbered Fig. 16 in Fieber’s key but is numbered both 16 and 17 on the plate (bottom left on each figure shown here); O. cuspidatus (not shown) is numbered Fig. 17 in Fieber’s key but is numbered 18 on the figure; O. apiculatus (not shown) is numbered Fig. 18 in Fieber’s key but is numbered 19 on the figure.

Fig. 5. Unpublished figures from FIEBER’s (1876) ‘The Cicadines of Europe’. a–b – Oliarus quinquecostatus; c – O. melanochaetus; d – O. limbatus.

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Appendix 6. Comments on synonymy in Oliarus by LÖW (1883: 147–148) (original and translation).

Fig. 6. Comments on synonymy in Oliarus by Löw (1883).

Translation of the comments on synonymy in Oliarus by LÖW (1883: 147–148)

Note: Added text is in square brackets.

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About the synonymy of the Cicadaria By Paul Löw in Vienna

Oliarus Panzeri m. (= leporinus Panz., Kirschb., Marsh., Scott, Fieb. et auct. ree. nec. Lin.) Oliarus leporinus Lin., Sahlbg. (= pallidus H.-Sch.)

J. Sahlberg, who certainly knows the cicades local to the home country of Linné [Sweden] better than any other author, in his “Öfversigt af Finlands och den Skandinaviska halföns Cicadariae” [1871] page 386 put the Flata pallida of Herrich- Schäffer [1835, locality not known] with Linné’s [1761] Cicada leporina and made the comment, that Linné was likely to have had Herrich-Schäffer’s species in front of him when he was describing his Cic. leporina, because his description does not fit any other species of the cixiids local to Sweden and that the species described by Herrich-Schäffer [1837] and Kirschbaum [1868] with the name of “leporina Lin.” has not been observed either in Scandinavia or in Finland and therefore surely is not Linné’s species.

Fieber [1876] did not consider these comments at all in his work “Les Cicadines d’Europe” listing the 01. pallidus H.-Sch. as a valid species, although he saw an identical, Swedish specimen of Cic. leporina Lin. in the collection of Germar. Fieber describes under the name of 01. leporinus Lin. a different species, namely the one that Panzer (Faun. Ins. Germ. 61, tab. 19) erroneously described and illustrated as Linné’s species, and which furthermore Kirschbaum [1868] (Cicad. v. Wiesbad, pag 45), Marshall [1864] (Ent. M. Mag. I. pag 155) and Scott [1870] (ibid. VII. pag 120) list by the name of leporina Lin.

The equivalence of 01. pallidus H.-Sch. and leporinus Lin. is also confirmed by M. Reuter (Meddel. Soc. pro F. et Fl. fenn. V. 1880, pag 195) through the interesting message, that 01. pallidus H.-Sch. is the only species in this genus which occurs in Scandinavia and Finland thus Linné could only have described this species as Cic. leporina in his “Fauna suecica”.

As therefore the name of Fieber’s [leporina] species needs to be changed, I think it would be suitable to name it after its discoverer Oliarus panzeri. The insect described and illustrated as Flata leporina Lin. by Herrich-Schäffer [1837] (Deutsch. Ins. 144, tab. 4) is neither Linné’s nor Panzer’s species but another, until now mysterious species”.

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Appendix 7. The description of Flata pallens Germar, 1821 (original and translation)

Fig. 7a. The original description of Flata pallens (from GERMAR 1821: 101–102). (Left.)

Fig. 7b. Type of Flata pallens Germar (ZML, Lviv). (Above.)

Translation of the original description of Flata pallens (from GERMAR 1821)

2) Flata pallens mine. Head obtuse; whitish; head and chest, black. Habitat in Tauria. Steven.

Slightly larger than F. nervosa, from the head to the tips of the elytra 4 par. lines long. The head at the front obtusely rounded – incidentally, entirely as in F. nervosa – black, all raised margins and keel yellow. The pronotum very short, cut in a sharp angled manner at the top in the middle, yellow, black in the recesses. The mesonotum yellow, rhomboidal, flat at the top in the middle, with five longitudinal keels. The tegulae at the base of the elytra, which all Fulgorids have, yellow white. The elytra again twice as long as the abdomen, dull yellowish white, yellow veins, small step-like tranverse veins, as the tips of all longitudinal veins, smoky grey. The wings unstained, the veins away from the last cleavage, black. The [p. 102] underside black, the edges of the rings of the abdomen yellow. The legs yellow, on the femur, a black long line.

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Appendix 8. The description of Cixius quinquecostatus Dufour, 1833 (original and translation) Note. Léon Jean Marie (or Jean-Marie Léon) Dufour (1780–1865) was a French medical doctor and naturalist whose collection of insects is preserved in MNHN. The following original description of R. quinquecostatus, appeared in an article on the anatomy and physiology of insects (DUFOUR 1833) and it is presumably for this reason that Dufour describes the digestive tract in some detail. A further description of the latter appeared some years later (DUFOUR 1839).

Fig. 8. The original description of Cixius quinquecostatus (from DUFOUR 1833: 224).

Translation of the original description of Cixius quinquecostatus from DUFOUR (1833) GENUS XXIV. – CIXIUS, CIXIE. SPECIES I. C.5-COSTATUS. New. Black, glabrous, back of thorax with 5 elevated lines, orbits of the eyes, prothoracic legs mottled with deep reddish brown; hemelytra grey, nerves finely brown spotted, abdominal segments with light reddish margins. Habitat in dry meadows. Length: 2 1/4 lines [= 0.5 cm approx.]. The Cixie with five ribs has all the structure and aspect of C. nerveuse, but is smaller than the latter and is very distinct from it as a species, either by the five projecting longitudinal lines on the rear of the thorax, or by the greyish colour of its hemelytra. I do not find it mentioned in the works of Entomology that are at my disposal. The digestive tract of the Cixius perfectly resembles, in structure and in configuration, that of the European Fulgora; so that the description of the one matches point by point that of the other. The hepatic vessels, of a sulphur yellow, very varicose and as articulated, number four, like in Fulgore but they present this characteristic, to some extent generic, that they meet two by two, before their insertion into the ventricle, in quite a distinct collar, rather long, diaphanous and smooth. 82 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Identity of some early described European Cixiidae

Appendix 9. Notes on the type series of Cixius quinquecostatus Dufour (= melanochaetus of authors) Type material examined. Syntypes: 1 ♂, Spain?; 1 ♀, no data; 1 specimen without abdomen and data (coll. Dufour, MNHN).

Remarks. In the original description of this species DUFOUR (1833) gave no details of the specimens he examined (see Appendix 8). However, the unlabelled specimen without abdomen could be the same that was dissected in order to give the description of the digestive system and later a more detailed description of the same (DUFOUR 1839). Later, Dufour says that he collected a male and female in copula in August 1829, which might be the same male and female in the collection (Figs 9a–d). Only this male has a data label, that could say (in an indistinct hand) the single word ‘Llayo’ (Cillorigo de Liébana), in the north of Spain, on the opposite side of the Pyrenees from Saint Sever, where Dufour lived. As these specimens do not disagree with the original description of C. quinquecostatus they are regarded as putative types. The following additional information (paraphrased) on Dufour’s collection has been supplied by Adeline Soulier-Perkins (MNHN): when Dufour died, Joseph Alexandre Laboulbène recurated the collection for the museum, but it was not in good condition. Perris reconditioned the specimens in new boxes with all the original labels at the bottom of the boxes. Some of the Dufour types stayed in the Edouard Perris collection that is in Montpellier. I have checked a copy of the catalogue of Perris’s collection but there is no mention of the Cixius quinquecostatus type, so the types should be the specimens in Paris.

Fig. 9. Syntypes of Cixius quinquecostatus Dufour (= R. melanochaetus auctt.). a–b – habitus male and female respectively; c–d – male genital segment, ventral view.

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Appendix 10. Identity of Reptalus limbatus (Fieber) The following figures of a male (no data, Puton coll., MNHN) are supplied by H. Hoch (see text). Two other specimens (Putton coll.) are also present: 1 ♂, no data; 1 ♀, ‘Ciudad Real’.

Fig. 10. Male genitalia of Reptalus limbatus (Fieber). a, b – pygophore; c–e, g–h – paramere; f – anal tube; i–j – aedeagus. Scale bars: 0.1 mm.

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Appendix 11. Key to genera of Cixiidae occurring in Britain Note. In the British Handbook to Fulgoromorpha (LE QUESNE 1960) the 12 species of Cixiidae were placed in two genera (Oliarus and Cixius). In the subsequent British checklist (LE QUESNE & PAYNE 1981) one of these species (C. pilosus (Olivier)) was placed in Tachycixius and C. borussicus Wagner was synonymised with C. cambricus China, and an additional species, Trigonocranus emmeae Fieber, was included. However, these species are currently placed in five genera following HOLZINGER et al. (2003), which can be separated in the following key. All male British specimens examined by us with five scutellar carinae and short head are here identified as R. quinquecostatus (second half of couplet 2). Similar females are assumed to be the same species. The only other cixiid with five scutellar carinae, Pentastiridius leporinus, has a longer head (compare Figs 11a and 11d).

1 Mesonotum with 5 carinae (Figs 11a, d–e)...... 2 – Mesonotum with 3 carinae...... 3 2 Vertex longer than broad (Fig. 11a); 1st and 2nd hind tarsomeres with more than 10 apical teeth (Fig. 11b); styles without long inner spine...... Pentastiridius (leporinus) – Vertex shorter than broad (Fig. 11d–e); 1st and 2nd hind tarsomeres with no more than 8 apical teeth (Fig. 11c); styles with long inner acute spine (Fig. 11f)...... Reptalus (quinquecostatus) 3 Outer apical margin of forewings with tubercles...... Tachycixius (pilosus) – Outer apical margin of forewings without tubercles...... 4 4 Forewings clear, without black or brown markings; size < 4 mm...... Trigonocranus (emmeae) – Forewings with black or brown markings; size > 6 mm...... Cixius (8 species)

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Fig. 11. Britsh Cixiidae. a–b – Pentastiridius leporinus, a – adult female (Paul Brock, 2011); b – apex of hind leg; c–d – Reptalus qinquecostatus auctt.: c – apex of hind leg; d – adult female (Tristan Bantock, 2011); e – adult male, Canvey Is. (BMNH); f – male genital capsule, caudal view.

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Appendix 12. Distribution and ecology of Reptalus quinquecostatus (of authors) The discovery that all British material identified as R. panzeri is the same as the R. quinquecostatus of Continental European authors is all the more surprising given that the latter has only been recorded from central and southern Europe (HOLZINGER et al. 2003) and while there are numerous British records, NICKEL (2003) notes that there are only six records for the whole of Germany, all confined to the northeast. R. quinquecostatus (of continental authors) is regarded as economically important as a potential vector of stolbur phytoplasma to both grapevines (PINZAUTI et al. 2008) and maize (BERTIN et al., 2010) in parts of central Europe, and so reliable identification is extremely important.

The following are the known continental records for R. quinquecostatus:

Austria: HOLZINGER (2009). Bulgaria: GJONOV (2004). Czech Republic: southern Moravia (DLABOLA 1956, LAUTERER 1957, MALENOVSKÝ et al. 2011, MALENOVSKÝ & LAUTERER 2012). France: 2 ♂♂ Rhone Valley, Chazay d’Azergues, vii.1995, W. della Giustina coll. (examined in the current study). Germany: confined to upper Rhine plain and Franconia; Speyer, vii.1989, Staffelbach, vii.1949, Coburg, vii. 1950, Kitzingen, vii.1994, Gerolzhofen, viii.1994, Erlangen, vii.2001 (NICKEL 2003). : in saltmarshes (NICKEL loc. cit.); in vineyards, Andornaktálya (BERTIN et al., 2010). Italy: widespread in sampled areas: Piemonte and Emilia Romagna (BERTIN et al., 2010). : in potato fields, Fundulea, Baragon Plain and Csikszereda (Miercurea- Ciuc) (BERTIN et al., 2010). : abundant in carrot fields, Baèka; record could refer to additional species (DROBNJAKOVIÆ et al. 2010). Slovakia: DLABOLA (1954). Slovenia: mostly on coastal shrubs and grass vegetation, vi. 2004, vii. 2001 and vii 2004 (SELJAK 2004). Turkey: LINNAVUORI (1965), DEMIR (2008). Ukraine: LOGVINENKO (1975); Azov, S. Ukraine (BERTIN et al. 2010).

In addition, BOURGOIN (2013) cites records of R. quinquecostatus from Spain, Portugal and Greece. The extra-limital range of R. quinquecostatus apparently extends through central Asia east to China (BOURGOIN 2013), however it has not been possible to verify the identity of specimens within Asia, and they are not included here, although it has recently been cited from Iran (MOZAFFARIAN & WILSON 2011). In contrast to continental Europe, the British records of R. quinquecostatus (as R. panzeri) are considerable. Also, all records are confined to southeast England, i.e. from Oxfordshire, Berkshire, Sussex, Surrey, Hampshire, the Isle of Wight, Essex and Kent (LE QUESNE 1960, KIRBY 1992) with the majority of records from Kent (KIRBY 1992). More recently, R. quinquecostatus has been recorded from a number of sites in south London (Beckenham, Dulwich, Crystal Palace, Windsor Great Park) and in Sussex (Bexhill-on-Sea, Westfield, Billingshurst, Newhaven) (JONES & HODGE 1999). At one of the London sites (Crystal Palace, in July 1996) very high population densities have been recorded, with up to several hundred specimens present (JONES & HODGE 1999).

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Recorded habitats for R. quinquecostatus in Britain include agricultural and fallow fields (DUFFIELD 1963, KIRBY 1992), urban parks, wood pasture, riverine meadows, brownfield sites (JONES & HODGE 1999) and coastal grazing levels (RAMSAY 2009). In contrast the habitat data for continental European localities is limited but in Germany R. quinquecostatus is associated with scattered scrub and tall herbs on sand or loamy soils, whilst in Hungary the species is recorded on saltmarsh (NICKEL 2003). Although recorded from a variety of habitats, the overriding habitat preferences of R. quinquecostatus are for heavier soils which periodically dry out and surface crack, whilst retaining their moisture. This is presumed to be the mechanism by which females are able to access plant roots in the soil on which to lay their eggs. Cixiidae nymphs are soil dwelling and feed on roots and potentially fungi (WILSON et al. 1994). DUFFIELD (1960) noted that R. quinquecostatus (as Oliarus panzeri) was widely distributed in East and West Kent (southeast England) and for many years was abundant in one field at Brook (Kent), where in August it could always be swept and beaten from sallow. Duffield went on to say that the field used to crack badly which he thought gave the insects a place to oviposit, and that ‘since the cracks have gone the insect has also gone, and has not been seen now for four years’. Similarly, KIRBY (2001) suggested grasslands in Britain that are wet in winter but dry out and crack in summer as the preferred habitat of R. quinquecostatus, and a recently collected specimen by Ramsay was from the margin of brackish ditches in grazing levels, dominated by Bolboschoenus maritimus (Sea Club-rush), 29 July 2008 at Stoke Marshes, Isle of Grain, north Kent (TQ8575), a habitat which also cracks and dries out in the summer months. Of interest is that the closely related Pentastiridius leporinus was also recorded from the same site, the only recorded co-occurrence of these two species in Britain, although they co-exist widely in continental Europe (e.g. BERTIN et al. 2010). Whether R. quinquecostatus requires woody plants to complete its development as in many other cixiids (WILSON et al. 1994), is unclear, but it seems that they are not essential for longer term population survival given that populations of R. quinquecostatus can persist in grassy fields with no other vegetation present (KIRBY 1992). Woody plants are often present in adjacent habitats and include Salix spp. (DUFFIELD 1960, EDWARDS 1896), Tamarix spp. (EDWARDS 1896) and parkland trees (JONES & HODGE 1999), and it is likely that the polyphagous adults are ‘vertical migrants’ (e.g. NICKEL 2003) onto taller hostplants, and have less exacting requirements than the nymphs.

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Appendix 13. The type concept FIEBER’s (1875) early comment on the ‘types’ he had examined (see Appendix 3b) prompted the following research on the subject of the first use of the word ‘type’ and how the type concept has changed over time. Much of the text is paraphrased from FARBER (1976) and VAN DER HAMMEN (1981). In the former work it is noted that the type concept during the first half of the nineteenth century was not a simple notion; it was used in at least three different ways: in classifications, collections and in morphology (see below). The word ‘type’ itself is derived from the Greek noun typos, which originally referred to a mould (a hollow form or matrix). The term was used by Plato (427–347 B.C.) to mean either an impression, a model or an outline or survey. Similarly, in the Lexicon Philisocophicum by Johannes Micraelius (1597–1658) the term typus is defined as (1) the original model of which any resemblance is made and (2) an example signifying something beforehand, i.e. a symbol. The word ‘archetype’ originally ‘arche’ was referred to by Aristotle (284–322 B.C.) in the Metaphysics as the beginning, starting-point, foundation, origin, cause, directing principle or ruler, all having in common to be the first from which something either exists, or comes into being, or becomes known. The word ‘type’ became replaced by ‘archetype’, as an example or the ‘original’ in the morphology type concept and was widely used in zoology (see ‘Visualizing the Archetype’ in WILLIAMS & EBACH 2008), reaching its pinnacle in the work of Richard Owen (1804–1892) in his development of the type-concept in comparative anatomy. The replacement of ‘type’ by ‘archetype’ could be connected with the introduction of ‘type’ in the rules for zoological nomenclature by the committee, appointed by the British Association for the Advancement of Science (see 1878 below), of which Owen was a member. The following are important dates in the use of the word ‘Type’ in the classification and collection type concept.

1775, 1778 Before the nineteenth century only the classification type concept was used, sometimes implicitly. For example, in BUFFON’s (1749–1789) ambitious project: a complete natural history of all the animals, in the sections on birds he firstly uses the word ‘type’ but latter does not, as follows:

1“Si l’on prend le rollier d’Europe pour type du genre…” “If we take the European Roller for type of the genus…” BUFFON (1775: 128). 2“Nous conserverons le nom générique de Gobe-mouche à celui d’Europe, comme étant généralement connu sous ce seul & même nom. D’ailleurs ce gobe-mouche nous servira de terme de comparaison pour toutes les autres espèces.”

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“We shall keep the generic name ‘Flycatcher’ for the European one, as it is generally known under this single name. Moreover, this flycatcher will serve for comparison with all other species’’. BUFFON (1778: 517–518).

1801, 1802, 1804 Buffon’s idea was developed by LAMARCK (1801) and LATREILLE (1802, 1804). The former stated the following:

“Pour faire connaître d’une manière certaine les genres dont je donne ici les caractères, j’ai cité sous chacun d’eux une espèce connue…. Et j’y ai joint quelques synonymes que je puis certifier; cela suffit à me faire entendre.” “In order to make known, without a doubt, the genera of which I give here the characters, I have cited under each of them a known species .... And I have added some synonyms that I can attest; this is enough to make myself understood”. LAMARCK (1801: page viii of the ‘Avertissement’).

LATREILLE (1802) mentioned under each of his genera one or more species that were indicated as examples (‘exemples’). On p. 64 of this work he characterized one of his examples moreover as the insect which had served him in the formation of the genus and later (LATREILLE 1804: 399) used the word ‘type’ (‘Le genre gamase a pour type la mite des coléoptères de Geoffroi…’). Evidently, Latreille’s types, as Buffon’s before him, were selected as standards for comparison (a comparison enabling the conception of a model of the genus) in such a way that now constitutes the basis of type-species selection in modern systematic zoology.

1826 The ‘idea’ of a type in the sense of an original described specimen appeared in the early nineteenth century, e.g., KIRBY & SPENCE (1826) gave the following advice when keying a specimen (added text in square brackets):

“When all these [efforts] fail, as they sometimes will, the dernier resort is a reference to the cabinet containing the original specimen from which the description was drawn…and thus many mistakes rectified, which would otherwise greatly mislead”. KIRBY & SPENCE (1826: 553)

1828 The fact that the word ‘type’ was not in frequent use at this time, is reflected in its absence in most dictionaries of the first half of the nineteenth century. A notable exception is Noah Webster, An American Dictionary of the English Language (New York:

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Converse, 1828), which gives this definition: ‘In natural history, a general form, such as is common to the species of a genus, or the individuals of a species.’

1837 The further development of the type-concept in systematic zoology is closely connected with the development of rules for zoological nomenclature. This development dates back to a paper by STRICKLAND (1837); although type-species are not mentioned, according to Strickland’s rule no. 18, families and subfamilies should be based on the most typical genus.

1838 The idea that a figure could serve the same purpose as the type specimen was demonstrated by BONAPARTE’s (1838) comments on the magnificent bird illustrations that appeared at this time:

“Throughout the list I have quoted as Types of the Species under consideration, the figures of the great works…” . BONAPARTE (1838: vi, preface).

1840 By 1840 there was near universal application of the classification type-concept as demonstrated by GRAY’s (1840) catalogue title: ‘A list of Genera of Birds, with an Indication of the Typical Species of Each Genus’ and in the same year, WHEWELL (1840), stated the following (added text in square brackets):

“A Type is an example of any class, for instance, a species of a genus, which is considered as eminently possessing the characters of the class” [In other words is typical of its group]. WHEWELL (1840: 494).

This is also an important date with regards to Westwood’s ‘Synopsis’ and all the type designations therein, which led to an ICZN Opinion that the phrase ‘typical species’ in WESTWOOD (1840) should be construed as a type designation of a genus: International Commission on Zoological Nomenclature (ICZN 1941).

1842 A committee was appointed by the British Association for the Advancement of Science in 1842, to consider the rules by which zoological nomenclature might be established on a uniform and permanent basis. Strickland, Darwin and Owen were among the members of this committee. In its rules, published in 1843, type-species of genera are mentioned but not type specimens and in the 1878 edition (p. 7–8), the type-species is connected with the typical portion of the original genus.

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1845 Similarly, the idea of ‘fixing’ a model or name carrier for a group was given by STRICKLAND (1845) in the following way: “every specimen is separately enumerated with its locality and the name of its donor, which is especially important in a collection containing the type-specimens, from which original descriptions have been made..” STRICKLAND (1845: 215).

“We may obtain a great amount of fixity, in the position at least, if not in the extent of our groups, by invariably selecting a type, to be permanently referred to as a standard of comparison..” STRICKLAND (1845: 219).

1845 Another early mention of a type specimen was by Adam White (Assistant in the Zoology Department of the British Museum) who commented on “Gray’s type specimen of P. Fiendii [Insecta: Coleoptera] from Mr. Children’s collection” in the BMNH. WHITE (1845: 110).

1850 In a series of catalogues (from 1850) of the British Museums insect collection the importance of the data of individual specimens was noted in John Gray’s Introductions. In those parts by Walker (e.g., WALKER’s 1850), individual specimens with their data are actually indicated by issuing each with a different letter, a, b, c etc. The use of a similar lettering system in other catalogue parts denoted only a difference of locality or collections. Walker’s system may have been the earliest enabling subsequent workers to know the actual number of specimens in the type series and hence, in the case of a single specimen, that what we now call a holotype had been designated.

1875 Further use of the word ‘type’ by FIEBER (1875) (see Appendix 3b under Dr Mayr, Dr Redtenbacher and Dr Stål).

1876 MCCOY (1876) is held to have designated a lectotype by referring to the cranium of a fossil marsupial as “the first described type of the species” (ICZN 1999: Article 74.6.1.2, under example).

1886 In the Code of Nomenclature adopted by the American Ornithologists’ Union (1886) rules are given for basing species and subspecies on ‘a type-specimen’.

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1888 Although some type categories are credited to Thomas (see 1893 below) he noted that the use of ‘co-type’ was from C. O. Waterhouse. Although Thomas gave no date or reference for Waterhouse his assertion is supported by the fact that some John Scott British Hemiptera type specimens in the BMNH, London, from 1888, bear a ‘co-type’ label and which, it is assumed, were placed on the specimens by a museum curator under the instructions of Waterhouse, who worked at the museum between 1866–1910.

1893 The modern concept of a primary type is credited to THOMAS (1893), who gave suggestions for a more definite use of types and restricted the term ‘type’ to a single specimen and also introduced the terms ‘paratype’ and ‘co-type’ (= syntype) (but see 1888 above).

1896 A review of the rules of nomenclature, including the use of types, as used by entomologists at the end of the nineteenth century, was published by WALSINGHAM & DURRANT (1896).

1897 The realisation of the importance of type specimens can be deduced from the type catalogues that appeared in the second half of the 19th century and which were commented on by Strickland (see 1845 above) and later by SCHUCHERT (1897) in the following way:

“All naturalists concede that type-specimens constitute the most important material in a museum of natural history. The true appreciation of this fact, however, is of recent date, and is shown in numerous lately published catalogues of types possessed by different museums…”. SCHUCHERT (1897: 636).

Summary The collection type-concept, although a technical concept, was not without its ambiguities. With regards to the type of a species, THOMAS (1893) summarised the situation, at that time, in the following way:

“The word ‘type’ itself when first introduced was meant to refer to the particular specimen (in the singular) originally described, but it soon was naturally applied to any individual of the original series, if more than one specimen was examined by the describer. In this there was little cause for confusion, but more recently it has been applied to any individual from the collection of the original author, obtained no matter

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how much later, and often not even determined by him as belonging to his species. Of late a still further cause of confusion has been introduced by certain authors who, obtaining specimens from the typical locality, have spoken of them as ‘typical specimens’, a method of reference which, although due to a praiseworthy regard for geographical exactness, is yet certainly liable to rise to inconvenience and confusion”.

With the growth of collections over time, taxonomists often had more than one specimen to use for descriptions of new species, and then could apply the word ‘type’ to more than one specimen. Conversely, sometimes a label saying ‘type’ was only put on one specimen when more than one specimen in the type series was present, while the word ‘Holotypes’ (for male and female) was used as late as 1930 by Pruthi in his material examined (PRUTHI 1930)! Despite the frequent occurrence of the word ‘type’ on labels, this did not always go hand in hand with its use in the description, thereby allowing for the subsequent designation of a lectotype.

Obviously, the word ‘type’ meant different things to different people, and in time an entire nomenclature grew up such that FRIZZEL (1933) was able to list 233 uses for the term ‘type’ while FERNALD (1939) added a few more usages and made the follow comment:

“The writer has been unable to learn who first completing his description attached to the specimen a label marked ‘Type’, Typical specimen or some other distinctive word or phrase. But whoever he was, he certainly raised the lid of Pandora’s box, beginning the liberation of a list of terms now over one hundred in number”.

The emergence of the collection type-concept was also made possible by two developments of late eighteenth and early nineteenth-century natural history. The first of these was the invention of reliable taxidermic procedures. These techniques were especially critical in ornithology, where specimens could not easily be preserved in good condition. For those who assembled the early collections the preservation of some specimens was an impossibility, as noted by RÉAUMUR (1747) in the following way:

“..ayant eu le déplaifir de les voir détruire journellement par des Infectes voraces, malgré les foins employez pour les défendre contre leurs dents..”

“I had the Mortification to see them every Day destroyed by ravenous Insects, in spite of all the care that had been taken to preserve them against their Teeth” (RÉAUMUR 1748: 305)

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By the early decades of the nineteenth century, however, the major problems had been solved and taxidermy had become part of the standard technical literature. The second development that permitted the collection type-concept to emerge was the growth and proliferation of museum and other large collections that were accessible to researchers. Institutions, such as the BMNH and the Paris Museum, served as repositories for the enormous numbers of specimens sent back to Europe by explorers, collectors, and naturalists in the first half of the nineteenth century.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 97–142, 2013

A review of the genus Macrocixius with descriptions of five new species (Hemiptera: Fulgoromorpha: Cixiidae)

ANDRÁS OROSZ Hungarian Natural History Museum, Baross u. 13, Budapest, H-1088, Hungary; e-mail: [email protected]

OROSZ A. 2013: A review of the genus Macrocixius with descriptions of five new species (Hemiptera: Fulgoromorpha: Cixiidae). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 97–142. – The genus Macrocixius Matsumura, 1914 (Hemiptera: Fulgoromorpha: Cixiidae) is reviewed. Two previously described species, M. giganteus Matsumura, 1914 and M. grossus Tsaur et Hsu, 1991, are redescribed, and five additional species are described as new: M. emeljanovi sp.nov. (from Taiwan and Nepal); M. gigantomimus sp.nov. (from Vietnam); M. lautereri sp.nov. (from Indonesia: Sumatra); M. monticola sp.nov. (from Vietnam); and M. orophilus sp.nov. (from Nepal). M. grossus and M. giganteus are recorded for the first time for Vietnam. All species are illustrated and keys for their identification are provided for the males. The morphology of the genitalia of females is discussed in detail with respect to diagnostic characters. Keywords. Auchenorrhyncha, planthoppers, taxonomy, new records, morphology, female genitalia, postfrontal area, Oriental Region

Introduction The genus Macrocixius Matsumura, 1914 was established by MATSUMURA (1914) for the newly described M. giganteus Matsumura, 1914, designated as type species. It was known at the time from Japan and Taiwan. TSAUR et al. (1991) redescribed the genus and the type species in detail with several new distributional records and described an additional species, M. grossus Tsaur et Hu, 1991. Based on extensive material collected mainly by the staff of the Hungarian Natural History Museum in the Oriental Region (largely in Nepal, Vietnam and Taiwan), the species of Macrocixius are here revised and five described as new for science. Identification keys are presented for the males, and the morphological diagnostic features of the females, including the female genitalia, are discussed in detail.

Material and methods The majority of the 125 specimens studied are deposited in the Hungarian Natural History Museum; a few specimens were loaned from other collections. The following abbreviations are used for the depositories of material mentioned in text:

97 A. OROSZ

HNHM ...... Hungarian Natural History Museum, Budapest, Hungary MMBC ...... Moravian Museum, Brno, Czech Republic NCHU ...... National Chung Hsing University, Taichung, Taiwan NMNS ...... National Museum of Natural Sciences, Taichung, Taiwan NTU ...... National Taiwan University, Taipei, Taiwan TARI ...... Taiwan Agricultural Research Institute, Wufeng, Taiwan ZIN . . . . . Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia

All the specimens in this project were allocated an individual code number (Nos. 1–125); these were also attached to the specimens. In order to distinguish them from the codes for collecting events, the individual code numbers appear in bold type in the text. Identification of all males involved study of the genitalia. Detached abdomens were boiled in 10% KOH solution in a water bath for 10–12 minutes and then rinsed in distilled water. After examination, the genitalia were preserved in glycerol in a plastic microvial, which was pinned with the specimen. Wings were largely studied and illustrated on slide mounts; any unique identifiers disclosed were also attached to the specimens. In females, after morphometric measurements had been taken, the abdomen was detached and transferred into distilled water, then photographs taken and drawings made of the anal tube and gonapophysis IX. The study of the internal female genitalia (vagina, spermatheca, bursa copulatrix) started with boiling the abdomen in 10% KOH solution and washing it in distilled water then, after the abdominal plates had been removed, the genital structures were stained with chlorazol black solution, after CARAYON (1969). In a couple of instances it proved necessary to relax the whole abdomen in distilled water (without boiling) and then, after removal of the abdominal plates, to stain it with eosin and treat it with chlorazol for the lumen of the bursa copulatrix and structures of the spermatheca to become accessible to photography (Figs 144, 148). Digital photographs of the genitalia were taken with a Leica MZ 95 or a Nikon Coolpix 4500.

Explanations and codes used for morphological terminology and morphometric measurements Measurements were taken from all specimens; data summarized for each species are given in Tables 1 and 2. Features measured and/or described were: 1. Postfrontal area (PFA). PFA is a trapezoid area bounded by the transverse carina at the apex of the vertex and the upper transversal carina of the frons (Fig. 10). Three measurements of the PFA were taken (Fig. 10: A, B and C) and two ratios calculated from them (A/B, C/D). 2. Vertex. Length of the vertex along midline (Fig. 10: D). 3. Frons. The ratio of the length (FRL) to the width (FRW) of the frons was established for males. FRL is here defined as the distance between the middle ocellus and the upper transversal carina, while FRW is the greatest distance between the two lateral carinae. 4. Stylus. Two measurements of the distal lobe were taken: diameter measured in the direction of the stem (DLL), and its width perpendicular to it (DLW) (Fig. 63). In dry preparations, the edges of the distal lobe were sometimes slightly rolled inwards or

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creased, so measurements of these may have been slightly inexact; however, this kind of error was considered negligible. The values of DLL/DLW ratio are given in the description of every species and in the key to the males. 5. Caudal margin of pygofer. The shape of the caudal margin of the pygofer in males (PCM) proved to be species-specific (Figs 69–76). 6. Aedeagus bears appendages, partly on both sides of the apex of the phallotheca, partly on either the apex of the flagellum or in its proximal third (Figs 11–13). The number and position of these appendages are diagnostic at species level. Codes are introduced for the appendages of the aedeagus in order to simplify the descriptions. In a ventral view of the phallotheca, lateral appendages on the right side are labelled as PHR1–4 (Fig. 13), and lateral appendages on the left side as PHL1–3 and, in the same aspect, appendages on the flagellum are labelled as FL1, FL2 and FL3 (Figs 11–12). 7. Ovipositor. Gonapophysis IX (GAPIX) bears on its dorsal surface a single, blunt, tooth-like process (clearly visible in lateral view: Fig. 9, see also Figs 145–147). This structure has been termed “middle tooth” in Cicadellidae by LINNAVUORI & AL NE’AMY (1983: Fig. 13). Also on the dorsal side of GAPIX in lateral view, there is a row of denticles extending to the middle of the GAPL section; the length of the row and the character of the teeth taken together appear to be a useful diagnostic feature at species level. The shape of GAPIX was studied in all female specimens; characteristic formations are illustrated in Figs 105–143 and 145–147. In many cases gonapophysis VIII adheres closely to the surface of gonapophysis IX, making it seem as if the lower, ventral side were also toothed (Figs 7–8); this can be quite confusing workers relatively unfamiliar with the group. The following morphometric characters were measured or calculated for GAPIX in females (Fig. 9): distance of the middle tooth from the apex (GAPL); width of GAPIX across the middle tooth (GAPW); GAPL/GAPW ratio; length of denticulate portion (TPRT) and GAPL/TPRT ratio; the last-named proved useful in identification of females to species level (Fig. 169). 8. Vagina in Cixiidae was described in general by BOURGOIN (1993); it consists of two parts: a posterior vagina and an anterior vagina (VA). A bursa copulatrix is attached to the dorsal part of VA, while a spermatheca and an antero-vaginal gland are attached to VA anteriorly. A common oviduct opens into the antero-ventral part of VA. Several plate-like sclerotized plates may be observed in the ventral and dorsal walls of VA; their pattern provides useful diagnostic information for the identification of females (“vagina wall pattern” VAGPT). 9. Spermatheca in Cixiidae was discussed by REMANE & ASCHE (1979) and BOURGOIN (1993); the latter author defined a diverticulum ductus (DD), longitudinally surrounded by a spiralling ductus receptaculi (DR, Figs 144, 148, 151–157), together forming a “double helix”. Diameters of DD (DDD) and DR (DRD) were measured and the number of the coils (THRS) of DR noted. The DRD values provided in Table 2 are averages calculated from measurements of all the coils in one specimen (Fig. 104). 10. Length of body was measured from apex of vertex to apex of forewings.

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11. Length of tegmen was measured from tegula to apex of forewing. 12. The anal tube has three defined parts: a stem, an apical lobe pointing towards the rear, and a short anal style (Fig. 15). In females, the ratio of the anal tube length/width as viewed from above (ANSL/ANSW) was measured and calculated.

Taxonomy Genus Macrocixius Matsumura, 1914

Macrocixius Matsumura, 1914: 393; TSAUR et al. (1991): 3; FENNAH (1956): 459; HUA (2000): 88. Type species: Macrocixius giganteus Matsumura, 1914, by original designation and monotypy. Redescription. The redescription of the genus given below is based on the original description (MATSUMURA 1914) and an extended redescription by TSAUR et al. (1991), complemented by some new descriptive notes. Size. Macrocixius includes the largest members of . For measurements of all species, see Tables 1–2. Colour. Body ventrally and dorsally yellowish-brown or brown, lightly covered in powdery wax; frons brightly brown, frontoclypeal suture with bright yellow spots near lateral carinae. Carinae of mesonotum yellowish-brown, brighter than basic colour; tegmina translucent, variably with brownish-black spots; veins yellowish-brown, occasionally with brown spots; labium yellowish-brown. Structure. Head between eyes narrower than pronotum. Vertex strongly concave, twice as wide as long along midline; apically with strong transverse carina, more or less continuous; median carina flattened apically, not reaching apex. Frons concave, as wide as long, widest at the level of antennae, apically tapering (see Table 1 for values of the FRL/FRW ratio), with well-developed lateral carinae, a median carina flattened apically and with transverse apical carina more or less straight. Between the transverse carinae of vertex and frons is a clearly-distinguishable, trapezoid postfrontal area; the A/B ratio of this area is a diagnostic feature for identification of certain species (see Fig. 168 and Tables 1 and 2). Pronotum tapered, with lateral carinae, medially strongly incised. Mesonotum tricarinate, medially protruding, descending laterally. Tegmina longer than abdomen, tectiform, veins with tubercles bearing setae; the two anal veins joining at clavus mid-point and running farther along the inner margin; twelve apical cells present. Hind wing with veins M3+4 and Cu1a separated, connected by a transverse vein. Hind tibia with 3(–4) lateral spines and a crown of 6 spines distally. Chaetotaxy of hind tarsi (7–)8/9. Labium three-segmented, its apex reaching the hind trochanter. Male genitalia. Pygofer structured as a symmetrical socket; its caudal margin (PCM), in lateral view, more or less rounded in a species-specific manner (Figs 69–72, 74–76). Apical lobe of anal tube with median and distal parts rounded and elongated towards the rear (Figs 14, 17, 19, 22), or medially slightly incised, with two lateral rounded projections (Figs 25, 28, 31), providing a diagnostic character for species groups. Stem of anal tube straight in lateral view (Figs 15, 23, 26) or sinuate (Figs 18, 20, 29, 32); in ventral view, elliptical (Fig. 30), nearly circular (Fig. 27) or narrow basally

100 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius and widening towards the apical lobe (Figs 16, 21, 24, 33, 62). Aedeagus with two parts: phallotheca and flagellum, the latter connected to the apical part of the phallotheca. Phallotheca with two basal triangular lobes (TRA) directed downwards (e.g. Figs 11, 35, 55) and spine-like appendages on both sides apically, near basal part of the flagellum: 2 or 4 on the right side and 2 or 3 on the left side (in ventral view). Flagellum also with 2 or 3 appendages (in left lateral view). The number of appendages, their position, size and orientation are diagnostic for each species. Genital styli composed of two parts: a stem, joining the connectivum, and a widening distal lobe (DL). The stem of the stylus slightly curved outwards in ventral view, with a distinct constriction midway along the inner side; both styli, in ventral view, thus together forming two sides of an oval. Distal lobes of styli turned outwards from axis; the width/length ratio (DLW/DLL, see Fig. 63) varies between species (see key to the males). Female genitalia. Pygofer nearly oval in caudal view, with two wax-producing plates fused medially. Anal tube nearly parallel-sided in dorsal view; for values of ANSL/ANSW ratio, see Table 2. GAPIX with a gentle protuberance and a row of small teeth; Table 2 for morphometric characters. Posterior vagina sac-like. Surface of bursa copulatrix with peculiar ornamented structure, composed of tiny pits including 2–7 setae (Figs 149, 150). Common oviduct opening into ventral part of anterior vagina (VA); walls of VA with characteristic sclerotized plates, their pattern (vagina wall pattern, VAGPT) an important diagnostic feature for individual species (Figs 77–103, 157–160). Spermatheca attaching to dorsal part of VA, laterally to antero-vaginal gland (AVG); spermatheca consisting of four clearly-distinguishable parts: ductus receptaculi (DR), followed by diverticulum ductus (DD), pars intermedialis, and glandula apicalis (Figs 144, 148, 151, 152: details, Figs 153–156, abbreviations in explanatory Fig. 104). Neither DRD/DDD ratio nor coil numbers (THRS) of DR proved species-specific (Table 2). Species included. The genus Macrocixius currently contains seven species, two of them described previously (M. giganteus and M. grossus) and five described as new in this paper (M. emeljanovi sp.nov., M. gigantomimus sp.nov., M. lautereri sp.nov., M. monticola sp.nov., and M. orophilus sp.nov.). Distribution. The genus is widespread in eastern and south-eastern Asia from Nepal to Sumatra, also known from parts of Japan, Taiwan, southern China and Vietnam. Biology, host plants and habitats. Information about the biology of Macrocixius spp. is quite scarce. The three Taiwanese species were collected between March and December, with most of them captured in September. Collecting data suggest a habitat preference from subtropical monsoon forests (M. grossus, M. emeljanovi: Taiwan, Kenting National Park, 120–200 m), hard-leaved rainforests (M. grossus, M. emeljanovi: Taiwan, Fu-Shan National Park ~600 m, Pingtung Co. ~ 900 m, Ming-Chyr 1180 m; Vietnam, Sa Pa 1650–1800 m, Da Lat 1500 m) to high-mountain mossy forests (M. emeljanovi, M. orophilus: Taiwan, Yakou, 2600 m, Hsiangyang 2200 m; Nepal, Ganesh Himal, 2720 m). There are no data on host plants, except for a general comment that a series of specimens representing M. grossus and M. emeljanovi was collected from the canopy in Taiwan (Wulu, 1100 m); all other material was collected at light traps.

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Key to species of Macrocixius based on external characters of males 1 Apical lobe of anal tube medially produced, rounded (Figs 14, 17, 19, 22) (giganteus species-group)...... 2 – Apical lobe of anal tube medially emarginate, concave (Figs 25, 28, 31) (grossus species-group)...... 5 2 Distal lobe of stylus relatively shorter (DLW/DLL >1.5; Fig. 66). Anal tube in ventral view V-shaped, apical part truncate (Fig. 62). Caudal margin of pygofer with a protruding, finger-like lobe (Fig.75)...... M. monticola sp.nov. – Distal lobe of stylus relatively longer (DLW/DLL <1.3). Anal tube in ventral view elliptical, apical part not truncate (Figs 16, 21, 24). Caudal margin of pygofer with no protruding lobe, truncate or rounded...... 3 3 Stem and apical lobe of anal tube, in lateral view, forming an obtuse angle (Fig. 15). Caudal margin of pygofer truncate in lateral view (Fig. 71)...... M. emeljanovi sp.nov. – Stem and apical lobe of anal tube, in lateral view, forming a near-right angle (Fig.20). Caudal margin of pygofer rounded in lateral view (Figs 70, 72)...... 4 4 Stylus with large protuberance on apical margin between distal lobe and stem (Fig. 67). Caudal margin of pygofer as Fig. 70...... M. giganteus Matsumura, 1914 – Stylus with small protuberance on apical margin between distal lobe and stem (Fig. 63). Caudal margin of pygofer as Fig. 72...... M. gigantomimus sp.nov. 5 Stylus with DLW/DLL ratio = 1.3. Anal tube, in ventral view, broadest in its apical third (Fig. 33); in lateral view, stem and apical lobe forming a right angle (Fig. 32), stem strongly sinuate. Pygofer with a pair of short lobes directed towards midline (PCM on Figs 73, 74). Chaetotaxy of first and second tarsomeres of hind legs: 9/9–11...... M. orophilus sp.nov. – Stylus with DLW/DLL ratio = 1.0. Anal tube, in ventral view, elliptical or rounded (Figs 27, 30); in lateral view, stem and apical lobe forming obtuse angle, stem tapering towards apical lobe or parallel-sided (Figs 26, 29). Pygofer with caudal margin rounded, lacking lobes. Chaetotaxy of first and second tarsomeres of hind legs: 7(–8)/9...... 6 6 Stem of anal tube elliptical in ventral view (Fig. 30), in lateral view tapering towards apical lobe (Fig. 29). Caudal margin of pygofer with a broadly rounded projection in lateral view (Fig. 69). Distal lobe of stylus elongate (Fig. 64)...... M. grossus Tsaur et Hsu, 1991 – Stem of anal tube almost circular in ventral view (Fig. 27); parallel- sided in lateral view (Fig. 26). Caudal margin of pygofer, in lateral

102 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

view, with a narrowly-rounded projection (slightly asymmetrical in the single specimen examined, Fig. 76). Distal lobe of stylus short, subtriangular (Fig. 58)...... M. lautereri sp.nov.

Key to species of Macrocixius based on characters of the aedeagus 1 Outline of basal third of phallotheca forming an obtuse angle in lateral view (Figs 34, 36)...... M. giganteus Matsumura, 1914 – Outline of basal third of phallotheca almost straight or very gently curved in lateral view (Figs 38, 40, 42, 44, 46, 48, 50, 52, 54, 56, 59, 60)...... 2 2 Flagellum with three appendages (FL1, FL2, FL3; Fig. 11), FL3 large, directed towards base of phallotheca (Figs 54, 56)...... M. grossus Tsaur et Hsu, 1991 – Flagellum with two appendages (FL3 absent)...... 3 3 Second appendage of flagellum (FL2) originating close to mid-point of flagellum (Figs 38, 46)...... 4 – Both appendages of flagellum originating near apex (Figs 42, 50, 59)...... 5 4 In left lateral view, appendage PHL2 adhering closely to side of flagellum, long and thin, appendage PHR4 sickle-shaped, curved beyond the stem of phallotheca and extending (in lateral view) over to the left half (Figs 46, 48)...... M. emeljanovi sp.nov. – In left lateral view, appendage PHL2 with ventro-caudal direction; PHR4 long, straight and directed mesiad (Figs 38, 40)...... M. monticola sp.nov. 5 Flagellum with appendages near apex differing in length, FL2 much shorter than FL1. Appendage PHR4 straight and thin, adhering closely to flagellum (Figs 59, 60)...... M. gigantomimus sp.nov. – Flagellum with appendages near apex of nearly equal length (Figs 42, 50). Appendage PHR4 absent (Fig. 52) or, if present, curved around stem (Figs 42, 44)...... 6 6 Phallotheca with two appendages on the right and two appendages on the left (Figs 50–52). Appendage PHR2 curved around stem to the left (Figs 50, 52); PHR4 absent. Orifice collar-like (Fig. 52, 53)...... M. orophilus sp.nov. – Phallotheca with four appendages on the right and two on the left (Figs 42, 44). Appendage PHR2 adhering closely to stem (Fig. 44); PHR4 present, curved around stem, extending over to the left. Orifice simple (Fig. 42)...... M. lautereri sp.nov.

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Identification of females Morphometric characters The only publication addressing M. giganteus and M. grossus (TSAUR et al. 1991), the two previously-described species, provides very little information about the females. The authors discriminated between the two species largely on the basis of characters of the male genitalia; for the females, they provided only drawings of the pygofer for the two species. In that contribution, the anal tube of M. giganteus is considered “very slender, much longer than wide” (length/width ratio about 3:1 in their Fig. 1J), while in M. grossus this is merely “slender” (length/width ratio about 2:1 in Fig. 2H; TSAUR et al. 1991). Calculations of this ratio (ANSL/ANSW) in a number of specimens identified as M. giganteus and M. grossus indicated that the ranges of intraspecific variability overlap partially in the two species (and in M. emeljanovi sp.nov. as well), rendering this character of only slight diagnostic value (Table 2). Altogether 42 females of Macrocixius spp. collected in Taiwan (33), Vietnam (7) and Nepal (2) were available for examination in this study (together with 83 males). The examination concentrated first on finding characters that might separate females of M. giganteus – M. grossus – M. emeljanovi from Taiwan; the results were then used to

Table 1. Measurements and ratios of Macrocixius males (in mm). See the Material and Methods section for explanations of character abbreviations.

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Table 2. Measurements and ratios of Macrocixius females (in mm). See the Material and Methods section for explanations of character abbreviations.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 105 A. OROSZ separate females of M. grossus from M. monticola. The following was disclosed in terms of morphometric characters: 1. Morphometric data for the postfrontal area and the vertex (A/B ratio) appear for all male and female specimens in Fig. 168. As the data for M. grossus encompass a wide range, females with values near low-point (No 83, 89, 94) needed to be checked by examination of other characters. One of the female specimens from Nepal (No 93) differed very distinctly from the other specimens (No 95, 96, 102) and fell into the range of M. emeljanovi. Subsequently, study of the genitalia confirmed the identity of this specimen as M. emeljanovi. A similar comparison was made with values of C/D ratio: they showed a similar distribution, but the differences were less convincing (see Tables 1–2 for values in males and females respectively). 2. A comparison of the length/width ratio of the frons (FRL/FRW) was made for most of the males. Among 30 M. emeljanovi males, thirteen M. grossus males from Taiwan and thirteen males from Vietnam, together with four males of M. monticola, none of the three species disclosed distinct, non-overlapping ranges (Table 1). This ratio is thus considered less than useful for separation of species of either sex. 3. The length/width ratio of the female anal tube (ANSL/ANSW) overlapped in M. giganteus (9 females measured), M. emeljanovi (23), and M. grossus (17). Only three specimens were available for M. monticola, probably too few to draw conclusions. 4. For gonapophysis IX, the GAPL/GAPW ratio was determined for 27 specimens, including six from Vietnam and 21 from Taiwan (Table 2). Because the data set was so limited, this ratio was not employed for species separation . 5. The GAPL/TPRT ratio was recorded in all female specimens and characteristic forms were illustrated with drawings and photos. It appears that this character is effective in discrimination between only a few species, e.g. M. emeljanovi from M. grossus in Taiwan, but it probably lacks such usefulness in the separation of certain continental Asian species.

Female internal genitalia The most reliable diagnostic characters for discrimination between Macrocixius spp. females may be found in the internal genitalia. Several specialists have addressed the internal female genitalia of the tribe Cixiini (REMANE & ASCHE 1979, BOURGOIN 1993, HOLZINGER 2002, HOLZINGER et al. 2002). BOURGOIN (1993) provided a description of the ectodermal genital ducts of Cixius nervosus (Linnaeus 1758), largely applicable to Macrocixius spp. as well. The descriptions that follow therefore draw much from that work. There are distinct sclerotized plates in the dorsal and ventral walls of the anterior vagina (vagina wall pattern, VAGPT). Since the bursa copulatrix and the spermatheca open dorsally, these sclerotized areas are best observed in ventral view (Figs 157–160, Figs 77–103); dorsal sclerites are indicated by a dotted contour in Figs 77–103. The sclerites of the vagina were studied in detail in M. giganteus (Figs 95–103, 159, 160), M. grossus (Figs 77–83, 157), and M. emeljanovi (Figs 86–94, 158) for which

106 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius several females were available from Taiwan. The number and shape of the sclerotized plates were shown to be species-specific in these three species and might presumably provide interspecific differences for other species of the genus as well. The spermatheca in Macrocixius has the same basic structure as that of Cixius Latreille, 1804, Tachycixius Wagner, 1939, Trirhacus Fieber, 1875, and others. A study of a large number of specimens of some species (particularly M. grossus, M. emeljanovi, and M. monticola) revealed that the structure of DR is quite variable between most species with respect to the diameters of the helices and the numbers of coils (THRS); these characters are therefore considered to be of no diagnostic value (Table 2).

Review of species of Macrocixius Macrocixius giganteus Matsumura, 1914 (Figs 12, 13, 19–21, 34–37, 67, 70, 95–103, 108, 134–143, 145–146, 156, 159, 160)

Macrocixius giganteus Matsumura, 1914: 394; TSAUR et al. (1991): 3; FENNAH (1956): 459; HUA (2000): 88. Syntypes: Japan, Kyushu, Kagoshima (coll. Hokkaido University Collection).

Material examined. Taiwan: 1 ♂, Taichung Co., Anmashan, 1650 m, 20.VI.1997, leg. B. Herczig & L. Ronkay (No 104, HNHM); 1 ♀, same data but 14.VII.1983, leg. C. T. Yang (No 108, NCHU); 1 ♂, Hualien Co., Tzuen, 5.IX.1986, leg. S. C. Tsaur (No 105, ZIN); 1 ♀, same data (No 110, NCHU); 1 ♀, Hualien Co., Lienhuachih, 5.IX.1988, leg. S. C. Tsaur (No 107, NCHU); 2 ♀♀, Hualien Co., Mukwa, 8.VIII.1985, leg. S. C. Tsaur (Nos 111 and 116, NCHU); 1 ♀, Ilan Co., Taipingshan, 4.VII.1988, leg. S. C. Tsaur (No 106, ZIN); 1 ♀, same data but 1.VII.1988 (No 114, NCHU); 1 ♀, Ilan Co., Chihtuan, 1.VIII.1986, leg. S. C. Tsaur (No 115, NCHU); 1 ♀, Chiayi Co., Tapan, 19.VI.1988, leg. S. C. Tsaur (No 109, NCHU); 1 ♀, Chiayi Co., Fenchifu, 12.VIII.1987, leg. C. T. Yang (No 113, NCHU). Vietnam: 1 ♀, No.285, Da Lat, 1500 m, 14–20.X.1988, leg. S. Mahunka & T. Vásárhelyi (No 94, HNHM).

Redescription. A detailed redescription has been given by TSAUR et al. (1991); what appears here is purely complementary. In TSAUR et al. (1991: 3), the captions to Figs 1 and 2 have inadvertently been transposed: Fig. 1 refers to M. giganteus and Fig. 2 to M. grossus. Colour. General colour yellowish-brown to brownish-black. Tegmen with yellow- to-green veins and several dark brown markings on veins and in cells. Male genitalia. Pygofer symmetrical; caudal margin (PCM) almost straight; entire outline of pygofer more or less sub-rectangular in lateral view (Fig. 70). Apical lobe of anal tube medially produced and rounded (Figs 19–21). Stylus with well-developed protuberance apically (Fig. 67); DLW/DLL = 1.26. Phallotheca, in lateral view, distinctly angled in basal third (Figs 34, 36), with four apical appendages near base of flagellum on right side (Figs 35–37), and with three appendages on the left surface. Flagellum near orifice with two acute, well-sclerotized appendages (Figs 34, 37). Female genitalia. The row of teeth at the apex of GAPIX is illustrated (in lateral view) in Figs 108, 134–143, 145 and 146. Anterior vagina, in ventral view, with three sclerotized plates in its ventral wall: one narrow and elongate, on right margin; the second well-defined, elliptical, located centrally; and the third and largest covering the caudal part (Figs 95–103, 159, 160). Spermatheca as in Fig. 156 (specimen No 94). Measurements and ratios in Tables 1–2 and Figs 168–169.

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Distribution. Japan (MATSUMURA 1914, MUIR 1925); China (FENNAH 1956, HUA 2000); Taiwan (SCHUMACHER 1915a, b; TSAUR et al. 1991, and additional records in the current study); Vietnam (first record in the current study, No 94). Habitat. Populations in Taiwan appear to inhabit various types of deciduous forests at altitudes of between 200 m and 2500 m. Differential diagnosis. Males of M. giganteus may be distinguished from all other congeners by apical lobe of anal tube medially produced, rounded (a defining feature of the giganteus species-group) and phallotheca distinctly angled in the basal third in lateral view (a unique character within Macrocixius). Females may be easily distinguished from other Macrocixius spp. by their characteristic vagina wall pattern (VAGPT), as in Figs 95–103 and 159–160. In distinguishing M. giganteus from the sympatric M. grossus, the GAPL/TPRT ratio is also diagnostic (Figs 108, 134–143, 145–146, 169, Table 2).

Macrocixius grossus Tsaur et Hsu, 1991 (Figs 3, 4, 11, 28–30, 54–57, 64, 69, 77–83, 104, 109–111, 112–123, 144, 148–150, 151–152, 154–155, 157) Macrocixius grossus Tsaur et Hsu, 1991: 3. Holotype: Taiwan, Taipei Co., Hsinhsien (coll. NTU).

Material examined. Taiwan: 7 ♂♂, 2 ♀♀, Ilan Co., Fu-Shan, 600 m, LTER site, 25–27.IX.2000, leg. L. Papp, L. Peregovits & L. Ronkay (Nos 1, 16, 61, 71, 73, 75–77, 84, HNHM); 6 ♂♂, 4 ♀♀ same locality, but 636 m, 15.IX.2007, leg. D. Rédei & J.-F. Tsai (Nos 66, 68–70, 74, 78–79, 81–82, 85, HNHM); 1 ♀, Lidao, 1240 m, 26.IX.2007, leg. D. Rédei & J.-F. Tsai (No 67, HNHM); 1 ♀, Taoyuan Co., Ming-Chyr, F.R.A., 1100 m, 5–6.X.1996, leg. Gy. Fábián & F. Nemes (No 15, HNHM); 1 ♀, Pingtung Co., on the road ‘199’, 900 m, 19.IV.1997, leg. A. Kun & L. Peregovits (No 88, HNHM); 4 ♀♀, Taichung Co., Tachen, 1890 m, 29.X.1996, leg. Gy. Fábián & F. Nemes (Nos 2–4, 7, HNHM); 2 ♀♀, Taichung Co., 14 km E of Puli, 430 m, 8.V.2010, leg. Gy. Fábián (Nos 117–118). Vietnam: 2 ♂♂, 1 ♀, Prov. Lao-Cai, Sa-Pa, 1650 m, 24.IX.1963, leg. T. Pócs (Nos 59, 65, 72, HNHM); 1 ♂, same, 10 km W of Sa-Pa, Fr. B. C., 1800 m, 17.III.1998, leg. L. Peregovits & T. Vásárhelyi (No 63, HNHM); 2 ♂♂, No.100, Moc Chau, 800 m, 25.X.1986, leg. T. Vásárhelyi (Nos 58, 60, HNHM); 1 ♂, 1 ♀, No.285, Da Lat, 1500 m, 14-20.X.1988, leg. S. Mahunka & T. Vásárhelyi (Nos 57, 89, HNHM); 1 ♂, 1 ♀, same but No.317, 17.X.1988 (No 64, 92, HNHM); 1 ♂, same but No.338, 19.X.1988 (No 56, HNHM); 1 ♂, same but No.302, 15.X.1988, leg. T. Vásárhelyi (No 62, HNHM). Description. Described in detail by TSAUR et al. (1991) in which. however, the captions for Figs 1 and 2 are inadvertently transposed (caption for M. grossus under the figure of M. giganteus and vice versa). The original description is complemented below. Colour. General colour brownish-black. Labium and bases of legs pale yellowish- brown. Tegmina with several black markings on veins and in cells, as Fig. 4. Male genitalia. Pygofer symmetrical, caudal margin (PCM) protruding medially in lateral view (Fig. 69). Apical lobe of anal tube medially emarginate in caudal view, concave, forming two small, rounded projections (Fig. 28); elliptical in ventral view (Fig. 30). Stylus symmetrical, distal lobe quite elongate-elliptical, DLW/DLL = 1.09 (Fig. 64). Phallotheca with four appendages apically on the right (Figs 55–57) and two appendages on the left. Flagellum with three appendages: FL3 well-developed, situated on the left in basal third, FL1 and FL2 acute and sclerotized, located near orifice (Figs 54, 57). Female genitalia. Ratios ANSL/ANSW and GAPL/GAPW as Table 2 and Fig. 169.

108 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

The range of genitalia between a number of females is shown in Figs 144, 148, 152 (from Taiwan), 151, and 157 (from Vietnam). Microsculpture of the bursa copulatrix as Figs 149 and 150. Spermatheca as Figs 154 and 155. Vagina wall pattern (VAGPT) of anterior vagina, in ventral view, with 4 or 5 sclerotized plates, rounded or elliptical, of variable size (Figs 77–82, 157), from which the small ventral-central plate is missing in some specimens (Figs 77, 80–81); the two adjacent anterior sclerotized plates may be fused (Figs 81–83). Measurements as Tables 1–2. Distribution. Taiwan (TSAUR et al. 1991; and further records in the current study); Vietnam (first record in the current study). Habitat. The specimens from Taiwan were collected at altitudes of 500–600 m in subtropical monsoon forest and hard-leaved rainforest. Most of the adults were taken in September and October. Specimens from Vietnam were collected by light-trapping in rainforest areas at altitudes of 800–1800 m. Differential diagnosis. M. grossus Males may be differentiated from other members of the genus by the medially emarginate, concave apical lobe of the anal tube (a character defining the grossus species-group) and the presence of a third appendage (FL3) of the flagellum (Fig. 54), while the other Macrocixius spp. have only two appendages. Females may be defined by high values of GAPL/TPRT (Fig. 169, Table 2), a short series of teeth on the GAPIX (Figs 109–123), and by a characteristic vagina wall pattern (VAGPT) composed of 4–5 small, rounded, sclerotized plates (Figs 77–82).

Macrocixius emeljanovi sp.nov. (Figs 5–9, 14–16, 46–49, 68, 71, 86–94, 124–132, 147, 158)

Type material. Holotype: ♂, “Taiwan, Ilan Hsien, Fu-Shan, LTER Site 600 m, 25–27.IX.2000. leg. L. Peregovits, L. Ronkay & L. Papp”/(No 23). Deposited in NMNS. Paratypes: Taiwan: 23 ♂♂, 6 ♀♀, same data as holotype (Nos 5–6, 12, 17, 19–20, 22–29, 32–34, 36–37, 39, 42, 44, 46, 52, 54–55; HNHM); 5 ♂♂, Yilan Co., Fushan B.G., 636 m, 16.IX.2007, leg. D. Rédei & J.-F. Tsai (Nos 38, 40–41, 43, 45; HNHM); 1 ♂, Taipei Co., Pi-Hu, 410 m, 26.III.1997, leg. G. Csorba & L. Ronkay (No 30; HNHM); 1 ♂, same data but 600 m, ca. 50 km SE of Taipei, 30.III.2000, leg. A. Kun & L. Peregovits (No 18; HNHM); 2 ♂♂, Pingtung Co., on the road ’199’, 500 m, 19.IV.1997, leg. A. Kun & L. Peregovits (No 35, 48; HNHM); 1 ♂, Taitung Co., Hsiangyang 2200 m, 13.VI.1997, leg. B. Herczig & L. Ronkay (No. 31; HNHM); 1 ♂, 1 ♀, Ilan Co., Fu-Shan B.G. LTER Site, 600 m, 25–28.III.2003, leg. M. Földvári & L. Papp (Nos 13, 47; HNHM); 1 ♂, 3 ♀♀, Taitung Co., Yakou, 2600 m, 6.XII.1998, leg. Gy. Fábián & Z. Korsós (Nos 9, 14, 21, 53; HNHM); 2 ♀♀, Pintung Co. Mutan, 12.VI.1997, leg. B. Herczig & L. Ronkay (Nos 10–11; HNHM); 1 ♀, Ilan Co. Ming-Chyr, Forest Recreation Area, 1200 m, 5.IV.2002, leg. Gy. Fábián & O. Merkl (No 8; HNHM); 1 ♀, Wulu, 1100 m, 26.IX.2007, leg. D. Rédei & J.-F. Tsai (No 87; HNHM); 1 ♂, 4 ♀♀, Kenting, 12.V.2010, leg. Gy. Fábián (Nos 119, 123, NCHU; No 122, ZIN; Nos 120-121, HNHM); 2 ♂♂, Taoyuan Co., Ming-Chyr, 1200 m, 4–5.V.2010, leg. Gy. Fábián (No 124, HNHM; No 125, ZIN); 1 ♀, Nantou, 8.VII.1975, leg. C. T. Yang (No 112, NCHU). Nepal: 1 ♀, Ganesh Himal, between Godlang-Nesim, 2720 m, 22.X.1995, leg. L. Peregovits & L. Ronkay (No 93, HNHM). Description. Colour. Ventral and lateral parts of body covered in scatterings of powdery wax of variable extent. Ground colour of body dark brown to blackish-brown; mesonotum the darkest part of dorsum, lateral carinae and area under and around tegulae

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 109 A. OROSZ paler, light yellowish-brown. Postfrontal area brown, marginal carinae with thin, brownish-black keel. Lateral portion of vertex near eye light yellowish-brown, central part darker, as pronotum; pronotum laterally dark brown behind compound eyes. Frons dark brown, central and lateral carinae brownish-black. Anteclypeus and postclypeus dark brown, labium pale brown. Abdominal dorsum and venter blackish-brown, posterior margin of sternites yellowish-brown. Trochanters, coxae and femora of fore- and mid- legs yellowish-brown. Ground colour of hind legs light yellowish-brown, femur and tibia with obscured brown spots. Tegmina translucent, main longitudinal veins with scattered blackish-brown tubercles, each provided with an apical seta, light yellowish-white in colour. Cross-veins at the tips of the wings and spots on membrane more or less blackish- brown (Fig. 6). Structure. Chaetotaxy of hind tarsi: 8/7(8). Male genitalia. Anal tube dilated from basal part to apical lobe (incrassate) in ventral view, nearly elliptical (Fig. 16); in lateral view, stem and apical lobe forming a slightly obtuse angle (Fig. 15); in posterior view, outline of middle and distal parts of apical lobe slightly produced and rounded (Fig. 14). Posterior margin of pygophore truncate (Fig. 71). Stylus as Fig. 68, DLW/DLL ratio approximately 1.3. Aedeagus as Figs 46–49. Basal part of phallotheca nearly symmetrical in ventral view; ventrocaudal outline of stem nearly straight in lateral view (Figs 46, 48), only gently curved from base to distal two- thirds; apex of phallotheca with pointed (spine- or blade-like) appendages on both sides; left side of stem with three appendages: pronounced PHL3 adhering closely to stem, and slender, spine-like PHL1 and PHL2 parallel to flagellum; right side of apex with four appendages, two of them large: a flattened, curved and sword-like PHR4 and a straight PHR3 directed downwards at acute angle to the stem. Flagellum with two strong, spine- like appendages (Figs 46, 49): FL1 smaller, located near orifice, FL2 larger, located in medial third of flagellum. Female genitalia. Vagina slightly asymmetrical in ventral view, left side slightly excavated midway; ventral wall with only two large sclerotized plates: anterior right plate with a characteristic shape, reminiscent of an outline of South America (Figs 86–94, 158), situated mid-vagina, tapering downwards towards its right side. Measurements in Tables 1–2 and Fig. 169. Etymology. The species is named after Professor A.F. Emeljanov, an eminent Russian entomologist. Distribution. The majority of the specimens collected to date are from Taiwan; single specimen from Nepal. Habitat. The altitude at which specimens were collected in Taiwan ranges between 120 m and 2600 m. The species was thus collected from lowland mixed broad-leave forest, through subtropical monsoon forest, deciduous hard-leaved forest at higher elevations, then subalpine coniferous forests, to high-mountain moss forest. Differential diagnosis. Males of M. emeljanovi sp.nov. may be distinguished from the other species of the genus by: apical lobe of anal tube medially produced, rounded (a character shared with other species of the giganteus species-group); stem of anal tube in

110 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius lateral view meeting the apical lobe at a slightly obtuse angle (Fig. 15); phallotheca, in lateral view, widening uniformly from basal third to apex (Fig. 46); PHR4 curved, sword- like, flattened in apical half (Figs 46, 48); FL2 arising from medial third of flagellum and parallel to it (Fig. 46). The females have a specific vagina wall pattern (VAGPT) with two large sclerotized plates (in ventral view), one of them of characteristic shape, reminiscent of South America and tapering towards the right (Figs 86–94, 158). The A/B ratios of both sexes are the lowest in the genus (Fig. 168). Remarks. From Nepal, only one female specimen (No 93) could be identified as M. emeljanovi based on the A/B ratio and a study of VAGPT, both of which match specimens from Taiwan (Fig. 90, compare Figs 86–94). This specimen from Nepal was collected in an undisturbed mossy forest area dominated by Quercus.

Macrocixius gigantomimus sp.nov. (Figs 22–24, 59–61, 63, 72)

Type material. Holotype: ♂, “Vietnam, Da Lat, Nr. 317, 1500 m, 17.X.1988. leg. S. Mahunka & T. Vásárhelyi”/(No 101). Deposited in HNHM. Paratype: 1 ♂, same data as holotype except Nr.285, 14.X.1988, leg. T. Vásárhelyi (No 103, HNHM). Description. Colour. General body colour lustreless dark brown. Frons between frontal suture and border ridge of postfrontal field blackish-brown, postfrontal area, middle of vertex, whole mesonotum and lateral sclerites under tegulae similarly coloured. Frontal suture separates dark areas of frons from -yellow postclypeus and anteclypeus, this yellow colouration extending towards genae; remainder of genae darker brown. Pronotum with two small, dark-brown pits close to midline, laterally light yellowish- brown. Genae covered with brilliant-white powdery wax on both sides below insertion of antennae. Legs yellowish-brown without conspicuous spots, apart from obscured darker spots on femora and tibiae near joints. Tarsal segments also darker, sparsely covered in powdery wax. Metatibia with 3–4 spines with dark tips laterally, following one another in a sparse row. Upper and lower part of abdomen honey-brown. Tegmina transparent, without traces of powdery wax, longitudinal veins yellowish-brown, except for scattered brown sections with tuberculi and dark brown setae originating from each tubercule; inner margin of tegmen with conspicuous brown spots at clavus midway; stigma dark brown; apical cells with scattered pale-brown spots; cross-veins blackish-brown; basal part of cubital vein with strongly thickened blackish-brown section. Structure. Chaetotaxy of hind tarsi: 8/9. Male genitalia. Caudal margin of pygofer (PCM) rounded in lateral view (Fig. 72). Stem of anal tube parallel-sided in lateral view, meeting apical lobe at a right angle (Fig. 23); stem elongate, elliptical in ventral view; in caudal view, middle part of apical lobe strongly produced and narrowly rounded (Figs 22, 24). Stylus, in inner dorso-lateral view, with a medially constricted stem; ratio DLW/DLL = 1.28, and with a small protruding knob on inner side apically between stem and distal lobe (Fig. 63). Phallotheca, in ventral view, parallel-sided in basal two-thirds, slightly narrowing

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 111 A. OROSZ distally; apex with spine-like appendages on both sides (Fig. 61): with three appendages in left lateral view (PHL1, PHL2, PHL3), PHL3 the largest, slightly curved to the left, PHL1 directed steeply towards inner body, and PHL2 adhering closely to stem, partly following the direction of PHL3 (Fig. 59); phallotheca with four appendages in right lateral view (PHR1–PHR4), PHR1 running parallel to PHL1, the other three appendages almost equal in size, slender, spine-like (Fig. 60). Flagellum with two appendages: a larger FL1, directed towards inner body, and a smaller FL2 originating on its side (Fig. 59). Female genitalia unknown. Measurements in Table 1. Etymology. The name refers to the close resemblance between M. gigantomimus sp.nov. and M. giganteus. Distribution. To date known from only Vietnam. Habitat. Captured with light traps at elevation of 1500 m, at the border of rainforest and cultivated fields. Differential diagnosis. The males of M. gigantomimus sp.nov. may be distinguished from all other congeners by the medially produced, rounded apical lobe of their anal tube (a character also shared with the other species of the giganteus species-group); anal tube parallel-sided in lateral view and forming a right angle between the stem and the apical lobe; PHR4 appendage of phallotheca long and straight, and smallest FL2 appendage of flagellum lying within the giganteus species-group.

Macrocixius lautereri sp.nov. (Figs 25–27, 42–45, 58, 76, 167)

Type material. Holotype: ♂, “W. Sumatra. Bengkulu prov. nr Curup; Bukit Kaba Mt.: 3°29′ S, 102°36′ E, 1000–1500m; D. Hauck leg.: 30.i.–3.ii.2000./Collectio Moravské Museum Brno/(No.100)”. Deposited in MMBC. Description. Colour. Vertex, pronotum and postfrontal area dark honey-coloured. Frons with irregular dark brown spots. Edges of middle and lateral carinae thinly brownish- black. Anteclypeus, postclypeus, and labium all yellowish-brown. Legs with similar coloration, partly with a darker brown tint. Mesonotum, in dorsal view, darker than remainder of body. Both dorsal and ventral surfaces of abdomen light honey-coloured. Tegmina translucent, covered in lightly-scattered powdery wax, main veins of tegmen yellow with tuberculi forming elongate brown spots on them. Apical crossveins blackish- brown (Fig. 167). Structure. Chaetotaxy of hind tarsi 7(8)/9; only second tarsomere with patellae. Male genitalia. Caudal margin of pygofer (PCM), in lateral view, forming a rounded lobe directed upwards (Fig. 76), sides slightly asymmetrical. Stem of anal tube nearly round in ventral view (Fig. 27); parallel-sided from base to apical lobe in lateral view, (Fig. 26), meeting apical lobe at almost right angles; apical lobe, in caudal view, emarginate midway, concave (Fig. 25), symmetrically rounded on sides. Stylus with

112 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius distal lobe forming a perfect equilateral triangle in dorso-lateral view (Fig. 58); DLL/DLW ratio 1:1. Phallotheca, in ventral view, with symmetrical downward-pointed spiny lobes on both sides basally (Fig. 43) and appendages on both sides apically: in left lateral view, with two appendages (PHL3, PHL1, Fig. 42): PHL3 with thickened basal part, oriented parallel to stem; in right lateral view, with four appendages (Fig. 44), of which PHR4 is the largest, flattened, wide, sword-like and curved around stem (in ventral view); other three appendages thin, elongate, spine-like. Flagellum with two short spine- like appendages (FL1, FL2), both situated apically, FL1 smaller, FL2 larger (Fig. 45). Female genitalia unknown. Measurements in Table 1. Etymology. The new species is named after Dr. Pavel Lauterer, entomologist emeritus of the Moravian Museum, with our great appreciation. Distribution. The only specimen known to date was found in Indonesia: Sumatra. Differential diagnosis. Macrocixius lautereri sp.nov. belongs to the grossus species- group in the light of the medially emarginate, concave apical lobe of its anal tube. It may be distinguished from both the other members of the group, M. grossus and M. orophilus sp.nov., by the shorter and evenly rounded apical half of its anal tube (in ventral view), and by the perfectly equilaterally triangular distal lobe of the style (Fig. 58). It also differs from M. grossus in the reduced number of appendages of the flagellum (only FL1 and FL2 present while M. grossus has three appendages); and from M. giganteus by the lack of the PHL2 appendage at the apical part of phallotheca (present in M. giganteus).

Macrocixius monticola sp.nov. (Figs 17, 18, 38–41, 62, 66, 75, 84, 105–107)

Type material. Holotype: ♂, “Vietnam, No.338, Da Lat. 19.X.1988. leg. S. Mahunka & T. Vásárhelyi”/(No 98). Deposited in HNHM. Paratypes. Vietnam: 1 ♂, same data as holotype but No.302, 15.X.1988. leg. Vásárhelyi (No. 2005-03-1003 [code of wing preparation], No 90); 1 ♂, Da Lat, 1500m, No.285, 14.X.1988. leg. Vásárhelyi T. (No 97); 1 ♂, Da Lat, Thac Prenn waterfall, 10.XII.1994. No.756, leg. S.Mahunka, Gy. Sziráki & L. Zombori (No 99); 1 ♂, Da Lat, Institute of Biology, 4.XII.1994, No.697, leg. S. Mahunka, Gy. Sziráki & L. Zombori (No 86); 1 ♀, Da Lat, Institute of Biology, 7.XII.1994, No.730, leg. S. Mahunka, Gy. Sziráki & L. Zombori (No 80); 1 ♀, Da Lat, 1500m, 14–20. X.1988, No.285, leg. S. Mahunka & T. Vásárhelyi (No 83); 1 ♀, Da Lat, No.358, 19.X.1988. leg. Vásárhelyi T. (No 91). All paratypes are deposited in HNHM. Description. Colour. Frons near frontoclypeal suture with pale yellowish-brown spots. Edges of median and lateral ridges of frons appearing as thin, dark-brown lines. Lateral parts of vertex near compound eyes largely yellowish-white. Pronotum and tegulae lighter than mesonotum, yellowish-brown. Metafemora with blackish brown spots at joints, remaining parts yellowish-brown. Forewings transparent, glassy, veins pale yellow, bearing brownish, longitudinally-elongated spots; tuberculi small, almost indistinguishable; in apical third of tegmen longitudinal veins and crossveins dark brown; symmetrical dark brown rows of spots on subcostal vein and inner margin of clavus. Structure. Chaetotaxy of hind tarsi: 8/8.

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Male genitalia. Pygofer symmetrical, caudal margin produced into finger-like lobe (Fig. 75). Anal tube, in ventral view, with stem constricted in basal quarter, widening in apical three quarters, V-shaped, with apex truncate (Fig. 62); apical lobe medially produced, slightly rounded (Fig. 17). Stylus axe-shaped in inner dorso-lateral view (Fig. 66); DLW/DLL = 1.64. Phallotheca narrowing towards flagellum (Fig. 39), apically with thin, pointed appendages on both sides: in right lateral view, with four appendages (PHR1–4, Fig. 40), of which PHR2 and PHR4 are larger and stronger and of nearly the same length, directed towards the inner body, PHR4 curved around stem; three appendages are visible near apex in left lateral view (Fig. 38), of which PHL2 is the longest, directed caudo-ventrad, and PHL3 is two-thirds as long as PHL2. Flagellum with strong, ventral, spine-like appendage FL2 (Fig. 38) and small FL1 near orifice. Female genitalia. Anal tube nearly parallel-sided in dorsal view. Serrate section of TPRT long, teeth distributed at regular intervals, large in mid-section, similar to those of M. giganteus. Sclerotized plates in wall of anterior vagina converging in ventral view and almost entirely filling the space between base and insertion point of ductus receptaculi (Fig. 84). Measurements appear in Tables 1–2 and Fig. 169. Etymology. The name of the new species is derived from the Latin monticola, indicating that it lives in mountains. Noun in apposition. Distribution. Currently known from only Vietnam. Habitat. Specimens of the type series were captured with light traps in two different habitats: at elevations of 900 m (near Thac Prenn waterfall) and around 1500 m (Da Lat) between arable land and rainforests. Differential diagnosis. M. monticola sp.nov. belongs to the giganteus species-group, based on the medially produced, rounded apical lobe of its male anal tube. The males of M. monticola sp.nov. differ from those of other members of this species-group in stylus with an axe-like distal lobe; V-shaped stem of anal tube, quite widely open, with truncate apex (in ventral view); symmetrical pygofer bearing finger-like process on its caudal margin; and flagellum of aedeagus with a strong and basally broad FL2, situated medio- ventrally (Fig. 38). The females may be distinguished from the other species of the genus by their specific vagina wall pattern (Fig. 84).

Macrocixius orophilus sp.nov. (Figs 1, 2, 31–33, 50–53, 65, 73, 74, 85, 133, 153)

Type material. Holotype ♂, “Nepal, Ganesh Himal, 2 km E of Thangjet, 2260 m, 1995.VII.20. leg. M. Hreblay & T. Csõvári”/(No 106). Deposited in HNHM. Paratypes: Nepal: 1 ♂, 1 ♀, same data as holotype (Nos 95, 96, HNHM). Description. Colour. General body colour dull blackish-brown. Frons, anteclypeus and postclypeus, postfrontal area and genae uniformly brown, lateral ridges yellowish-brown, margins thinly lined in black. Vertex, pronotum and mesonotum dark brown dorsally, mesonotum with lateral ridges lighter brown. Lateral parts of vertex near compound eyes

114 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius dull honey-brown. Hind legs yellowish-brown, only tips of tarsal segments darker, blackish-brown. Forewings translucent, longitudinal veins yellowish-white, bearing many longitudinally elongated brown spots, each with a tubercle; crossveins and stigma dark brown; apical cells spotted brown; costal vein and inner margin of tegmen thinly blackish-brown, both with a row of brown spots along their length (Fig. 2). Hind wing as Fig. 1. Structure. Chaetotaxy of tarsi: 9/(9–11);with one or two patellae on second tarsal segment. Male genitalia. Caudal margin of pygofer with a narrowly rounded lobe medially, curved towards inner body (PCM in Figs 73–74). Anal tube, in lateral view, with stem and apical lobe forming a near-right angle, ventral margin of stem sinuate, strongly bulging outwards medially; apical lobe, in caudal view, medially concave emarginate, with lateral parts rounded and protruding (Figs 31–33). Stylus, in dorso-lateral view, with stem parallel-sided, DLW/DLL = 1.33. Phallotheca asymmetrical in ventral view, stem slightly shifted leftwards at mid-section (Fig. 51); phallotheca apically with spine-like appendages on both sides: in left lateral view, with two appendages (PHL1, PHL2, Fig. 50), of which PHL 2 is larger, slightly sickle-shaped, directed towards inner body; in right lateral view, with two appendages (PHR1, PHR2, Fig. 52), of which PHR2 is larger, sword-like, curved around stem (in ventral view) to left. Flagellum with two spine-like appendages (FL1, FL2), both originating close to each other, near orifice. Orifice surrounded by characteristic collar-like structure (Figs 52–53). Female genitalia. The single female specimen examined lacks an anal tube. GAPIX with a row of small, uneven teeth near apex (Fig. 133). Sclerotized plates in wall of anterior vagina (VAGPT, Fig. 85) displaying a specific pattern, with most of the sclerotized plates located at basal (posterior) part on both sides. Measurements in Tables 1–2 and Fig. 169. Distribution. The species is currently known only from Nepal. Etymology. The name is derived from the Greek “ο′ρος” (oros = mountain) plus the Latinized Greek philus (= liking). Habitat. The type series was captured with light traps in high-mountain moss-forest. Differential diagnosis. M. orophilus sp.nov. belongs to the grossus species-group, having a medially emarginate, concave apical lobe of the anal tube. It may easily be distinguished from the other species of the genus by the genitalia of both sexes and by the specific chaetotaxy (9/9–11). Unique features of the male genitalia include the collar- like apical structure of the flagellum near the orifice (Figs 52–53) and the 2-2 apical appendages of the phallotheca (all the other species have totals of more than four appendages); the females have a characteristic vagina wall pattern (Fig. 85).

Acknowledgements First of all, I am greatly obliged to Igor Malenovský (Moravian Museum, Brno) for his useful advice on the manuscript and the loan of material; then to A. F. Emeljanov and V. N. Gnezdilov (Zoological Institute of the Russian Academy of Sciences, St.

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Petersburg, Russia), and Man-Miao Yang (National Chung Hsing University, Taichung, Taiwan) for loans of Macrocixius specimens held in their institutions. I am also grateful to S.-C. Tsaur (TARI Wufeng), who kindly sent me the basic literature on the Fulgoromorpha of Taiwan. For critical comments on the manuscript and for taking the photographs that accompany this article I thank my colleagues Eszter Lazányi, Dávid Rédei and Zoltán György; the help of László Ronkay during the preparation of the manuscript is also gratefully acknowledged.

References

BOURGOIN T. 1993: Female genitalia in Hemiptera Fulgoromorpha, morphological and phylogenetic data. Annales de la Société Entomologique de France (N. S.) 29(3): 225–244. CARAYON J. 1969: Emploi du noir chlorazol en anatomie microscopique des insectes. Annales de la Société entomologique de France (N.S.) 5: 179–193. FENNAH R. G. 1956: Fulgoroidea from Southern China. Proceedings of the California Academy of Sciences 28(13): 441–527. HOCH H. & REMANE R. 1985: Evolution und Speziation der Zikaden-Gattung Hyalesthes Signoret, 1865 (Homoptera, Auchenorrhyncha, Fulgoroidea, Cixiidae). Marburger Entomologische Publikationen 2(2): 1–427. HOLZINGER W. E. 2002: A review of the European planthopper genus Trirhacus and related taxa, with a key to the genera of European Cixiidae (Hemiptera: Fulgoromorpha). European Journal of Entomology 99: 373–398. HOLZINGER W. E., EMELJANOV A. F. & KAMMERLANDER I. 2002: The family Cixiidae Spinola 1839 (Hemiptera: Fulgoromorpha)–a Review. In: HOLZINGER, W. E. (ed.): Zikaden Leafhoppers, Planthoppers and Cicadas (Insecta: Hemiptera: Auchenorrhyncha). Denisia 4: 113–138. HUA L. Z. 2000: List of Chinese insects.Vol I. Zhongshan University Press. Guangzhou, 2+7+448 pp. LINNAVUORI R. E. & AL-NE’AMY K. T. 1983: Revision of the African Cicadellidae (subfamily Selenocephalinae) (Homoptera, Auchenorrhyncha). Acta Zoologica Fennica 168: 10–27. MATSUDA R. 1976: The internal female genitalia. Pp. 296–298. In: MATSUDA R.: Morphology and evolution of the insect abdomen. Pergamon Press, Oxford–New York–Toronto–Sydney–Paris–Frankfurt, 534 pp. MATSUMURA S. 1914: Die Cixiinen Japans. Annotationes Zoologicae Japonenses 8(3): 393–434. MUIR F. 1925: On the genera of Cixiidae, and (Fulgoroidea, Homoptera). The Pan- Pacific Entomologist 1: 97–110. OSSIANNILSSON F. 1978: The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark. Part 1: Introduction, infraorder Fulgoromorpha. Fauna Entomologica Scandinavica 7(1), Scandinavian Science Press, Kopenhagen, pp. 1–222. REMANE R. & ASCHE M. 1979: Evolution und Speziation der Gattung Cixius Latreille, 1804 (Homoptera Auchenorrhyncha Fulgoromorpha Cixiidae) auf den Azorischen Inseln. Marburger Entomologische Publikationen 1: 1–264. SCHUMACHER F. 1915a: Der gegenwärtige Stand unserer Kenntnis von der Homopteren-fauna der Insel Formosa unter besonderer Berücksichtigung von Sauter’schem Material. Mitteilungen aus dem Zoologischen Museum in Berlin 8: 73–134. SCHUMACHER F. 1915b: Homoptera. In: H. Sauter’s Formosa-Ausbeute. Supplementa Entomologica 4: 108–142. TSAUR S. C., HSU T. C. & VAN STALLE J. 1991: Cixiidae of Taiwan, Part V. Cixiini except Cixius. Journal of Taiwan Museum 44(1): 1–8.

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Figs 1–6. Wings of Macrocixius spp. 1–2 – M. orophilus sp.nov.; 3–4 – M. grossus Tsaur et Hsu: 5–6 – M. emeljanovi sp.nov. 1, 3, 5 – hind wing, 2, 4, 6 – tegmen. Scale bars = 3 mm (Figs 1, 3, 5); 5 mm (Figs 2, 4, 6).

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Figs 7–13. Morphology of Macrocixius spp. 7–9 – gonapophyses of M. emeljanovi sp.nov., in lateral view (No 93). 7 – gonapophyses VIII and IX together; 8 – gonapophysis VIII; 9 – gonapophysis IX. 10 – M. grossus Tsaur et Hsu, morphometrical characters of head (frons, postfrontal area, vertex). 11–12 – aedeagus in left lateral view, with terminology of appendices: 11 – M. grossus; 12 – M. giganteus Matsumura; 13 – M. giganteus, aedeagus in right lateral view, Abbreviations: A = width of upper carina of frons; B = median lenght of postfrontal area; C = width of vertex between the eyes; D = median lenght of vertex; FL1, FL2, FL3 = first, second and third appendages of flagellum; GAPL = lenght of gonapophysis IX from the middle tooth to apex; GAPW = width of gonapophysis IX; PFA = postfrontal area; PHL1, PHL2, PHL3 = first, second and third appendages of phallotheca on left side; PHR1, PHR2, PHR3, PHR4 = first, second, third and fourth appendages of phallotheca on right side; TPRT= lenght of the serrate part of gonapophysis IX; TRA = basal triangular lobe of phallotheca. 118 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 14–24. Male anal tube of Macrocixius spp. 14–16 – M. emeljanovi sp.nov. (No 93). 17–18 – M. monticola sp.nov. (No 90). 19–21 – M. giganteus Matsumura (No 105). 22–24 – M. gigantomimus sp.nov. (No 103). 14, 17, 19, 22 – apical lobe, caudal view; 15, 18, 20, 23 – anal tube, lateral view; 16, 21, 24 – anal tube, ventral view. Scale bars = 0.3 mm (Figs 14, 17, 19, 22); 0.5 mm (Figs 15, 16, 18, 20, 21, 23, 24).

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Figs 25–33. Male anal tube of Macrocixius spp. 25–27 – M. lautereri sp.nov. (No 100). 28–30 – M. grossus Tsaur et Hsu (No 73). 31–33 – M. orophilus sp.nov. (No 96). 25, 28, 31 – apical lobe, caudal view; 26, 29, 32 – anal tube, lateral view; 27, 30, 33 – anal tube, ventral view; 28, 31 – apical lobe, caudal view. Scale bars = 0.3 mm (Figs 25, 28, 31); 0.5 mm (Figs 26, 27, 29, 30, 32, 33).

120 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 34–37. Aedeagus of Macrocixius giganteus Matsumura (No 105): 34 – left lateral view; 35 – ventral view; 36 – right lateral view; 37 – flagellum, dorsal view. Scale bars = 0.5 mm.

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Figs 38–41. Aedeagus of Macrocixius monticola sp.nov. (No 90): 38 – left lateral view; 39 – ventral view; 40 – right lateral view; 41 – flagellum, dorsal view. Scale bars = 0.5 mm.

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Figs 42–45. Aedeagus of Macrocixius lautereri sp.nov. (No 100): 42 – left lateral view; 43 – ventral view; 44 – right lateral view; 45 – flagellum, dorsal view. Scale bars = 0.5 mm.

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Figs 46–49. Aedeagus of Macrocixius emeljanovi sp.nov. (No 93): 46 – left lateral view; 47 – ventral view; 48 – right lateral view; 49 – flagellum, dorsal view. Scale bars = 0.5 mm.

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Figs 50–53. Aedeagus of Macrocixius orophilus sp.nov. (No 96): 50 – left lateral view; 51 – ventral view; 52 – right lateral view; 53 – flagellum, dorsal view. Scale bars = 0.5 mm.

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Figs 54–57. Aedeagus of Macrocixius grossus Tsaur et Hsu (No 73): 54 – left lateral view; 55 – ventral view; 56 – right lateral view; 57 – flagellum, dorsal view. Scale bars = 0.5 mm.

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Figs 58–62. Male genitalia of Macrocixius spp. 58 – M. lautereri sp.nov., style, dorso-lateral view, inner face. 59–61 – M. gigantomimus sp.nov., aedeagus: 59 – left lateral view; 60 – right lateral view; 61 – ventral view. 62 – Macrocixius monticola sp.nov., male anal tube, ventral view. Scale bars = 0.5 mm.

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Figs 63–72. Genital styles and male pygofer of Macrocixius spp. 63, 72 – M. gigantomimus sp.nov.; 64, 69 – M. grossus Tsaur et Hsu; 65 – M. orophilus sp.nov.; 66 – M. monticola sp.nov.; 67, 70 – M. giganteus Matsumura; 68, 71 – M. emeljanovi sp.nov. 63–68 –style, dorso-lateral view, inner face; 69–72 – pygofer, lateral view. Abbreviations: PCM = caudal margin of pygofer. Scale bars = 0.5 mm.

128 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 73–85. Male pygofer and vagina wall patterns of Macrocixius spp. 73, 74, 85 – M. orophilus sp.nov. (85 – No 95): 75, 84 – M. monticola sp.nov. (84 – No 91); 76 – M. lautereri sp.nov., 77–85 – M. grossus Tsaur et Hsu (77 – No 92, 78 – No 7, 79 – No 67, 80 – No 89, 81 – No 70, 82 – No 15, 83 – No 118). 73–76 – pygofer: 73 – dorsal view; 74 – lateral view. 77 –85 – vagina wall patterns, ventral view. Abbreviations: PCM = caudal margin of pygofer. Scale bars = 1 mm (Figs 73–76); 0.5 mm (Figs 77–85).

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Figs 86–94. Vagina wall patterns of Macrocixius emeljanovi sp.nov. (several specimens, in ventral view: 86 – No 13, 87 – No 6, 88 – No 11, 89 – No 112, 90 – No 93, 91 – No 120, 92 – No 14, 93 – No 12, 94 – No 10). Scale bars = 0.5 mm.

130 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 95–103. Vagina wall patterns of Macrocixius giganteus Matsumura (several specimens, in ventral view: 95 – No 116, 96 – No 108, 97 – No 114, 98 – No 109, 99 – No 94, 100 – No 115, 101 – No 113, 102 – No 111, 103 – No 110). Scale bars = 0.5 mm.

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Figs 104–111. 104 – Macrocixius grossus Tsaur et Hsu, female inner genitalia, schematic drawing after Fig. 144 (No 66). 105–111 – Gonapophysis IX, in lateral view (in different species and specimens of Macrocixius collected in Vietnam): 105–107 – M. monticola sp.nov. (105 – No 80, 106 – No 83, 107 – No 91); 108 – M. giganteus Matsumura (No 94); 109–111 – M. grossus Tsaur et Hsu (109 – No 72, 110 – No 89, 111 – No 92). Abbreviations: BC = bursa copulatrix; DD = diverticulum ductus; DDD = diameter of the coil of diverticulum ductus; DR = ductus receptaculi; DRD1 = diameter of the first coil of ductus receptaculi; DRDN = diameter of the last coil of ductus receptaculi; GA = glandula apicalis; PI = pars intermedialis; VA = anterior vagina. Scale bar = 0.5 mm (Figs 105–111).

132 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 112–123. Macrocixius grossus Tsaur et Hsu, gonapophysis IX, in lateral view, in different specimens collected in Taiwan (112 – No 3, 113 – No 2, 114 – No 4, 115 – No 7, 116 – No 15, 117 – No 16, 118 – No 66, 119 – No 67, 120 – No 68, 121 – No 69, 122 – No 70, 123 – No 88). Scale bars = 0.5 mm.

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Figs 124–133. Macrocixius spp., gonapophysis IX, in lateral view. 127–132 – M. emeljanovi sp.nov. (several specimens: 124 – No 6, 125 – No 8, 126 – No 10, 127 – No 11, 128 – No 14, 129 – No 21, 130 – No 93, 131 – No 20, 132 – No 112). 133 – M. orophilus sp.nov. (No 95). Scale bars = 0.5 mm.

134 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 134–143. Macrocixius giganteus Matsumura, gonapophysis IX, in lateral view (several specimens: 134 – No 113, 135 – No 110, 136 – No 116, 137 – No 111, 138 – No 114, 139 – No 115, 140 – No 109, 141 – No 107, 142 – No 106, 143 – No 108). Scale bars = 0.5 mm.

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Figs 144–147. Macrocixius spp., female genitalia. 144–145 – Macrocixius grossus Tsaur et Hsu: 144 – female internal genital structures (No 66), 145 – gonapophysis VIII and IX together, lateral view. 146 – Macrocixius giganteus Matsumura, gonapophysis IX; lateral view (No 93). 147 – M. emeljanovi sp.nov., same (No 13).

136 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Review of the genus Macrocixius

Figs 148–150. Macrocixius grossus Tsaur et Hsu, internal female genitalia. 148 – internal genital structures of No 68. 149–150 – microsculpture of the bursa copulatrix (150 – enlarged part of Fig. 149, both No 15).

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Figs 151–152. Macrocixius grossus Tsaur et Hsu, internal female genitalia (151 – No 92, 152 – No 2).

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Figs 153–156. Macrocixius spp., spermatheca. 153 – M. orophilus sp.nov. (No 95); 154–155 – M. grossus Tsaur et Hsu (No 15, No 69); 156 – M. giganteus Matsumura (No 94).

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Figs 157–160. Macrocixius spp., internal female genitalia. 157 – M. grossus Tsaur et Hsu, internal genital structures (No 72). 158 – M. emeljanovi sp.nov., vagina wall pattern (No 13). 159–160 – M. giganteus Matsumura, same (159 – No 110; 160 – No 113).

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Figs 161–167. Macrocixius spp., habitus in dorsal view. 161–162 – M. emeljanovi sp.nov., males (161 – No. 119; 162 – No. 124); 163 – M. emeljanovi sp.nov., female (No. 121); 164 – M. giganteus Matsumura, female; 165–166 – M. grossus Tsaur et Hsu, females (165 – No. 1, 166 – No. 89); 167 – M. lautereri sp.nov., holotype male (No. 100).

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Fig. 168. A/B ratio (see Material and Methods and Fig. 10 for explanation) of Macrocixius specimens (horizontal axis, see the text for explanations of code numbers of specimens), grouped according to collecting localities (vertical axis): TW 1, 2, 4, 5 = Taiwan, Fushan Bot. Garden; TW 3, 6–9 = Taiwan, various localities and dates; VIET 1– 4 = Vietnam, various localities and dates; NEPAL = Nepal, various localities and dates. Symbols: rhombus = males, reversed triangle = females; different colours are used for different species: M. emeljanovi sp.nov. (red), M. giganteus Matsumura (black), M. gigantomimus sp.nov. (green), M. grossus Tsaur et Hsu (blue), M. monticola sp.nov. (ochre), M. orophilus sp.nov. (yellow). Arrows show specimens reidentified based on A/B values.

Fig. 169. GAPL/TPRT ratio (see Material and Methods section and Fig. 9 for explanation) of Macrocixius female specimens (horizontal axis), different species shown separately (vertical axis). Specimens of M. grossus and M. giganteus collected in Taiwan and Vietnam are marked with different symbols.

142 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 143–153, 2013

A new genus, Loisirella, and two new species of Bennarellini from Ecuador (Hemiptera: Auchenorrhyncha: Fulgoromorpha: Cixiidae)

WERNER E. HOLZINGER, INGRID HOLZINGER & JOHANNA EGGER Ökoteam-Institute for Animal Ecology and Landscape Planning, Bergmanngasse 22, 8010 Graz, Austria; e-mail: [email protected], [email protected], [email protected]

HOLZINGER W. E., HOLZINGER I. & EGGER J. 2013: A new genus, Losirella, and two new species of Bennarellini from Ecuador (Hemiptera: Auchenorrhyncha: Fulgoromorpha: Cixiidae). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 143–153. – A new genus, Loisirella gen.nov., and two new species, Loisirella erwini sp.nov. and Noabennarella paveli sp.nov., of the Neotropical clade Bennarellini are described from the Yasuní National Park in eastern Ecuador. Loisirella is the only Bennarellini genus with only two instead of five large sensory pits on its lateral abdominal appendages. A key to the genera and species of Bennarellini is provided. Keywords. Planthoppers, taxonomy, new genus, new species, Neotropical Region, Amazonian rainforest, terra firme forest, Yasuni National Park

Introduction The tribe Bennarellini Emeljanov, 1989 is a small Neotropical clade within the planthopper family Cixiidae (EMELJANOV 1989). Only three genera and four species have been described: Bennarella Muir, 1930 with B. bicoloripennis Muir, 1930 (known from Brazil and Guyana) and B. fusca Muir, 1930 (Brazil), and the two monotypic genera Amazobenna Penny, 1980 with A. reticulata Penny, 1980 (Brazil) and Noabennarella Holzinger et Kunz, 2006 with N. costaricensis Holzinger et Kunz, 2006 (Costa Rica) (MUIR 1930, PENNY 1980, HOLZINGER & KUNZ 2006). Two more taxa from Peru are mentioned in literature, but not formally described: an “undescribed genus” with very peculiar sensory pits is mentioned by HOCH (1987), and an undescribed Noabennarella species is figured by HOLZINGER & KUNZ (2006). Bennarellini are easily recognisable by the presence of unique abdominal appendages that are an autapomorphy of this clade (EMELJANOV 1989, HOLZINGER & KUNZ 2006). In the Cixiidae samples Terry Erwin and his colleagues collected in the Yasuní National Park in Ecuador, we recognised two more yet undescribed taxa from Bennarellini. Here we provide descriptions of these species, one of them also representing a new genus.

Material and methods The specimens were collected in the Yasuní National Park in eastern Ecuador (e.g. BASS et al. 2010). Both sampling sites are located in a terra firme Amazonian lowland forest: The first site is the Onkone Gare Station transect (ONK) in the Reserva Etnica

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Waorani (00°39′10″S, 76°26′00″W, 250 m) near the Piraña field station, the second one is in a distance of about 35 km from the first site close to the Tiputini Biodiversity station (TIP, 00°39′25″S, 76°27′10″W, 230 m). The insects were collected in 1994–1999 by the method of “canopy fogging” in a sophisticated sampling design that is described by LUCKY et al. (2002) and ERWIN et al. (2005). The Fulgoromorpha specimens were sorted out and preliminarily identified by Charles Bartlett and Lawrence Barringer (BARRINGER 2011). The holotypes designated here and most other specimens studied are stored in trust for the country of Ecuador at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (USNM). Five specimens of each species are kept in the collection of the Ökoteam-Institute for Animal Ecology and Landscape planning, Graz, Austria (OEKO), and two paratypes of Noabennarella paveli sp.nov. in the Moravian Museum, Brno, Czech Republic (MMBC).

Taxonomy Noabennarella paveli sp.nov. (Figs 1–13)

Type material. Holotype: ♂, “681 Ecuador Orellana TransectEnt.1km 220m S. Onkonegare Camp Reserva Etnica Waorani / 00°39′10″S 076°26′00″W 20.vi.94 T.Erwin et al fogging terra firme forest / Morphospecies Cixiidae 4 L. Barringer UDEL m.s.thesis / 2062”; stored in trust for the country of Ecuador at the National Museum of Natural History, Smithsonian Institution (USNM). Paratypes: Same locality as holotype (ONK): 1 ♂ 1 ♀, 4.ii.1996 (USNM); 1 ♂, 29.vi.1994 (OEKO); 2 ♂♂, 4.x.1994; 2 ♂♂ 1 ♀, 6.x.1994 (OEKO, USNM); 3 ♂♂ 1 ♀, 9.x.1994; 1 ♂ 1 ♀, 10.x.1994; 1 ♂ 1 ♀, 9.ii.1995; 9 ♂♂ 2 ♀♀, 1.vii.1995 (USNM, OEKO); 1 ♂, 3.vii.1995; 2 ♂♂, 6.vii.1995; 1 ♀, 4.x.1995 (OEKO); 2 ♂♂, 6.x.1995; 1 ♀, 4.ii.1996; 1 ♂, 13.ii.1996; 3 ♂♂, 23.vi.1996 (USNM, MMBC). Locality “TIP”: 2 ♂♂, 1.vii.1998; 1 ♂ 1 ♀, 4.vii.1998 (USNM, MMBC); 2 ♂♂, 22.x.1998; 2 ♂♂ 1 ♀, 7.ii.1999 (USNM). Description. Size. Medium-sized Cixiidae, body length in males 4.0–4.6 mm (6.5–7.5 mm including wings), in females 4.3–4.8 mm (7.4–7.8 mm including wings). Coloration. Head, body and legs straw-coloured, dorsally yellowish-brownish. Lateral keels of frons with large black spots (Figs 1, 2), reaching from lateral ocellus to epistomal suture. Fore wings (Figs 1, 10) in the basal half with a brownish tinge and brownish-yellow veins. Pterostigma whitish. Apical half of fore wings transparent with yellowish-whitish veins. Spots with a distinct brownish tinge present along a half-circular line from the distal end of the pterostigma along the small crossveins to the distal end of the clavus. Apical cells in some specimens distally with a brownish tinge, too. Hind wings colourless, transparent. Male genitalia. Shaft of aedeagus close to its base with two slightly curved, basally triangular spines, and three spines at the apex of the shaft: one long, almost straight, movable spine on the left side, and two tiny spines at the apex. Flagellum widening apically and bearing one straight spine on its inner left side (Figs 12, 13). Genital styles spoon-shaped, with a distinct inner lobe and a larger lateral lobe, the latter being produced into a long tip (Fig. 11). Anal segment very characteristic because of its long ventral process (Figs 6–8).

144 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genus and species of Bennarellini

Figs 1–5. Noabennarella paveli sp.nov. 1 – habitus of holotype, lateral view; 2 – head, ventral view; 3 – head, frontal view; 4 – head and prothorax, dorsal view; 5 – apex of female abdomen, lateral view.

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Figs 6–9. Noabennarella paveli sp.nov. 6 – male genital and anal segments (aedeagus and styli omitted), left lateral view; 7 – male anal segment, dorsal view; 8 – same, caudal view; 9 – ventro-median process of male genital segment, ventral view.

Female genitalia. Ovipositor (Fig. 5) and female genitalia very similar to those of the (still undescribed) Noabennarella species figured by HOLZINGER & KUNZ (2006: 60). Differential diagnosis. Morphology of N. paveli sp.nov. fits very well to the generic description of Noabennarella (HOLZINGER & KUNZ 2006), thus we consider it congeneric with the type species, N. costaricensis Holzinger et Kunz, 2006, and do not repeat the general description of the genus here. The new species is larger than N. costaricensis and differs also in coloration and characters of the male genitalia: In N. paveli, the black spots on the lateral keels of the frons are much larger than in N. costaricensis, reaching from the epistomal suture to the lateral ocelli (Fig. 1), the spines of the aedeagus are short and triangular in N. paveli (Figs 12, 13), but long and slender in N. costaricensis, and the anal segment of males of N. paveli has a large, distally widening ventral process (Figs 6–8), whereas this process is small and tongue-shaped in N. costaricensis. Etymology. The species is named in honour of our teacher and friend RNDr. Pavel Lauterer, a very well known Auchenorrhyncha and Psylloidea specialist.

146 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genus and species of Bennarellini

Figs 10–13. Noabennarella paveli sp.nov. 10 – fore wing; 11 – male genital style, inner maximum view; 12 – aedeagus, left lateral view; 13 – same, right lateral view.

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Loisirella gen.nov. Type species. Loisirella erwini sp.nov. Description. Small cixiids (body length 2.5–3.5 mm, including wings less than 6 mm) with laterally compressed body and wings in resting position steeply inclined. Frons almost twice as long as broad, concave, separated from (very small) vertex by a distinct and straight carina. Lateral carinae of frons strongly produced, median carina missing. Median and lateral ocelli distinct. Scapus short, ring-like, pedicellus almost spherical. Pronotum short, lateral carinae distinct. Mesonotum with indistinct median and lateral keels. Fore wings long (almost 3 times as long as wide), apically widening (Fig. 23), R and MA trifurcate, MP bifurcate. Metatibiae without macrosetae, apically with six spines. Metatarsus with 6+5 apical spines, without platellae. Abdominal segments 4 and 5 bearing lateral processes with two large sensory pits (Fig. 29). Male genital and anal segments almost symmetrical, aedeagus with shaft and flagellum well-developed. Apex of female abdomen of the “plesiomorphic type” within Cixiidae (see HOLZINGER et al. 2002), like in other Bennarellini: truncate, without wax plate, ovipositor evenly curved, adjacent to abdomen (Fig 30). Differential diagnosis. Loisirella gen.nov. is the only Bennarellini genus with abdominal appendages bearing only two large sensory pits. One sensory pit is present at the tip of the appendage of the fourth, the other one on the appendage of the fifth abdominal segment. The abdominal appendages of all other Bennarellini bear five sensory pits: the appendage of the fourth segment has three and the process of the fifth segment two apical sensory pits. The size of these pits in Noabennarella and Bennarella is about half of the size of Loisirella’s pits. Etymology. The genus is dedicated to “Mother Fulgoromorpha” Dr. Lois B. O’Brien. The name itself is an arbitrary combination of letters. Gender feminine. Note. The placement of Loisirella gen.nov. into the tribe Bennarellini is based on the fact, that its abdominal appendages are formed by the fourth and fifth abdominal segment and bear sensory pits with large setae. Thus we think that these appendages are homologous to the appendages of the hitherto known Bennarellini species and we consider Loisirella to be part of a monophyletic Bennarellini clade (possibly derived from a Oecleini-like Cixiidae taxon), characterised by these abdominal processes as autapomorphy. Species of another Cixiidae tribe, the palaeotropical Bennini Metcalf, 1938, (recently revised by HOCH 2013) also have lateral abdominal processes, but these are derived from the third and fourth abdominal segments and the shape of these appendages (see HOCH 2013: 18 ff. and HOLZINGER et al. 2002: 123, Fig 10E) is completely different to those of Loisirella. The only other taxon with lateral abdominal appendages within Fulgoromorpha is the family Achilixiidae, with again morphologically very different appendages (see WILSON 1989: 490 and HOLZINGER & KUNZ 2006: 56). Furthermore, based on characters of male and female genitalia, head, wing venation etc., Loisirella is not an Achilixiidae but a true Cixiidae s. str.

148 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genus and species of Bennarellini

Figs 14–17. Loisirella erwini sp.nov. 14 – habitus of holotype, lateral view; 15 – head, ventral view; 16 – head, frontal view; 17 – head and prothorax, dorsal view.

If we consider Loisirella as a member of Bennarellini, the question raises, if the configuration of two large sensory pits in Loisirella (instead of 3+2 in all other Bennarellini) represents a plesiomorphic state within Bennarellini or is a secondary reduction (by either loss or fusion of some pits). More detailed studies on the morphology of these appendages are needed to solve this question.

Loisirella erwini sp.nov. (Figs 14–25, 29, 30)

Type material. Holotype: ♂, “1572 Ecuador Orellana ErwinTransect Onkonegare Camp Reserva Etnica Waorani / 00°39′25.7″S 076°27′10.8″W 22.vi.96 T.Erwin et al fogging terra firme forest /Morphospecies Cixiidae 16 L. Barringer UDEL m.s.thesis / 2130”; stored in trust for the country of Ecuador at the National Museum of Natural History, Smithsonian Institution (USNM).

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Figs 18–25. Loisirella erwini sp.nov. 18 – male genital and anal segments (aedeagus and styli omitted), left lateral view; 19 – ventro-median process of male genital segment, ventral view; 20 – male genital style, inner maximum view; 21 – male anal segment, dorsal view; 22 – same, dorso-caudal view; 23 – fore wing; 24 – aedeagus, left lateral view; 25 – same, right lateral view.

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Figs 26–30. 26 – Noabennarella sp., abdominal appendage, ventro-lateral view. 27 – Bennarella bicoloripennis Muir, head and thorax in dorsal view; 28 – Noabennarella costaricensis Holzinger et Kunz, head in lateral view (from HOLZINGER & KUNZ 2006, slightly modified); 29 – Loisirella erwini sp.nov., lateral abdominal appendage; 30 – Loisirella erwini sp.nov., apex of female abdomen, laterocaudal view.

Paratypes: Same locality as holotype (ONK): 2 ♀♀, 22.vi.1996; 1 ♂ 2 ♀♀, 29.vi.1994; 1 ♂, 6.x.1994; 2 ♀♀, 9.x.1994; 2 ♂♂, 8.ii.1995; 1 ♀, 12.ii.1995; 1 ♂ 1 ♀, 6.vii.1995; 1 ♂, 6.x.1995; 2 ♂♂ 1 ♀, 8.x.1995 (all USNM); 3 ♂♂ 2 ♀♀, 7.ii.1996; 4 ♂♂ 3 ♀♀, 10.ii.1996; 4 ♂♂ 2 ♀♀, 2.x.1996 (USNM, OEKO). Locality “TIP”: 1 ♂, 5.ii.1999 (OEKO); 1 ♂, 6.ii.1999 (USNM). Description. Size. Small Cixiidae, body length in males 2.5–3.2 mm (4.8–5.7 mm including wings), in females 2.9–3.3 mm (5.0–5.8 mm including wings). Coloration. Head, body and legs yellowish-brownish, without distinct colour pattern. Fore wings with a brownish tinge and yellowish-brownish veins. Apical part of fore wings slightly darker in central part, only inner parts of apical cells almost colourless. Morphology of head, thorax and abdomen as in generic description (Figs 14–17). Male genitalia. Aedeagus with shaft slightly bent, without prominent keels and without spines. Flagellum with two slender spines: a long one emerging at the base of the

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 151 W. E. HOLZINGER ET AL. flagellum, and a short one emerging subapically (Figs 24, 25). Genital styles apically widened, bent inwards (Fig. 20). Anal segment more or less symmetrical, ventrally prolonged, tongue-like (Figs 21, 22). Etymology. The species is named in honour of Dr. Terry L. Erwin, famous rainforest ecologist and Carabidae specialist, who was the head of the team that collected this remarkable species.

Key to genera and species of Bennarellini 1 Lateral abdominal appendages apically with two large sensory pits (Figs 14, 29)...... Loisirella erwini sp.nov. – Lateral abdominal appendages apically with five large sensory pits (grouped 3+2; Figs 1, 26)...... 2 2 Median keel on frons distinct, elevated. Lateral keels of frons strongly elevated apically, semicircular in lateral view (Figs 2–4)...... 3. (Noabennarella) – Median keel on frons absent or vanishing. Lateral keels of frons less produced, directed obliquely laterad (Fig. 27)...... 4 3 Black spots on lateral keels of frons small, not reaching lateral ocelli (Fig. 28). Spines on the shaft of the aedeagus very long and slender. Male anal segment with a small tongue-shaped ventral process...... Noabennarella costaricensis Holzinger et Kunz – Black spots on lateral keels of frons large, reaching from epistomal suture to lateral ocelli (Fig. 1). Spines on the shaft of the aedeagus short, basally triangular (Figs 12, 13). Male anal segment with a large, distally widening ventral process (Figs 5–7)...... Noabennarella paveli sp.nov. 4 Vertex small, but present, separated from frons by a transverse carina. Cubital veins of fore wing entering anal margin at acute angle. First cell of cubitus nearly triangular, only twice as long as wide...... 5. (Bennarella) – Vertex not recognisable, as there is no subapical transverse carina on dorsal border of frons. Cubital veins entering anal margin at almost right angle. First cell of cubitus approximately ten times as long as wide...... Amazobenna reticulata Penny 5 Fore wing fuscous basally, translucent apically...... Bennarella bicoloripennis Muir – Fore wing uniformly dark fuscous...... Bennarella fusca Muir

Acknowledgements We are grateful to Terry Erwin, Washington, for enabling us to study the material he has collected, and for valuable information on the sampling sites. We thank Charles

152 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genus and species of Bennarellini

Bartlett and Lawrence Barringer, both Delaware, for their help with the material and additional support in our work, Igor Malenovský for his patience with us, Lois O’Brien for information on American Cixiidae, and Hannelore Hoch for valuable comments on the manuscript.

Zusammenfassung Eine neue Gattung und zwei neue Arten des neotropischen Tribus Bennarellini werden beschrieben: Loisirella gen.nov. mit Loisirella erwini sp.nov. sowie Noabennarella paveli sp.nov. Zudem wird ein Bestimmunggschlüssel zu den Gattungen und Arten der Bennarellini präsentiert. Loisirella ist die einzige Gattung der Bennarellini mit nur zwei an Stelle von fünf großen Sinnesgruben an den Enden der Lateralanhänge des Abdomens. Die Tiere stammen aus dem Yasuní Nationalpark im Osten von Ecuador und wurden im Rahmen eines umfassenden „Canopy fogging“-Forschungsprojekts gesammelt.

References

BARRINGER L. E. 2011: Canopy assemblages and species richness of planthoppers (Hemiptera: Fulgoroidea) in the Ecuadorian Amazon. Unpublished M.Sc. thesis, University of Delaware, Newark. BASS M. S., FINER M., JENKINS C. N., KREFT H., CISNEROS-HEREDIA D. F., MCCRACKEN S. F., PITMAN N. C. A., ENGLISH P. H., SWING K., VILLA G., DI FIORE A., VOIGT C. C., KUNZ T. H. 2010: Global conservation significance of Ecuador’s Yasuní National Park. PLoS ONE 5(1): e8767. doi.10.1371/journal.pone. 0008767, pp. 1–22. EMELJANOV A. F. 1989: To the problem of division of the family Cixiidae (Homoptera, Cicadina). Entomologicheskoe Obozrenie 68(1): 93–106 (in Russian, English translation published in Entomological Review 68(4): 54–67). ERWIN T. L., PIMIENTA M. C., MURILLO E. & ASCHERO V. 2005: Mapping patterns of α-diversity for across the western Amazon Basin: A preliminary case for improving conservation strategies. Proceedings of the California Academy of Sciences 56, Suppl. 1(7): 72–85. HOCH H. 1987: The tribe Bennini – a monophyletic group within the Cixiidae? Pp. 55–57. In: Arzone A. & Vidano C. (eds.): Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, 7–11 Sept. 1987. Consiglio Nazionale delle Ricerche; Turin, 652 pp. HOCH H. 2013: Diversity and Evolution of the South-East-Asian planthopper taxon Bennini (Hemiptera, Cixiidae). Nova Supplementa Entomologica 23: 1-296. HOLZINGER W. E., EMELJANOV A. F. & KAMMERLANDER I. 2002: The family Cixiidae Spinola 1839 (Hemiptera: Fulgoromorpha) – a review. Denisia 4: 113–138. HOLZINGER W. E. & KUNZ G. 2006: A new genus and species of Bennarellini form Costa Rica (Hemiptera: Fulgoromorpha: Cixiidae). Zootaxa 1353: 53–61. LUCKY A., ERWIN T. L. & WITMAN J. D. 2002: Temporal and spatial diversity and distribution of arboreal Carabidae (Coleoptera) in a western Amazonian rain forest. Biotropica 34: 376–386. MUIR F. 1930: Three new species of American Cixiidae (Fulgoroidea, Homoptera). Pan-Pacific Entomologist 7/1: 12–14. PENNY N. D. 1980: A revision of American Bennini (Hemiptera: Fulgoroidea: Cixiidae). Acta Amazonica 10(1): 207–212. WILSON M. R. 1989: The planthopper family Achilixiidae (Homoptera, Fulgoroidea): a synopsis with a revision of the genus Achilixius. Systematic Entomology 14: 487–506.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 155–162, 2013

Acoustic communication in the subtroglophile planthopper Trigonocranus emmeae Fieber, 1876 (Hemiptera: Fulgoromorpha: Cixiidae: Oecleini)

HANNELORE HOCH *, ROLAND MÜHLETHALER & ANDREAS WESSEL Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt- Universität zu Berlin, Invalidenstrasse 43, D-10115 Berlin, Germany; e-mail: [email protected], [email protected], [email protected] *Corresponding author

HOCH H., MÜHLETHALER R. & WESSEL A. 2013: Acoustic communication in the subtroglophile planthopper Trigonocranus emmeae Fieber, 1876 (Hemiptera: Fulgoromorpha: Cixiidae: Oecleini). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 155–162. – A recently discovered, and apparently autochthonous, population of Trigonocranus emmeae Fieber, 1876 in Germany (Rhineland-Palatinae: Bernkastel) confirmed the large degree of intraspecific variation with regard to the size of the compound eyes, length of tegmina and bodily pigmentation reported in the literature. For the first time, intraspecific communication signals for the species could be recorded. T. emmeae is identified to be at least subtroglophile, and its potential as a suitable model to study the evolution of cave adaptation in a natural system is discussed. Keywords. Auchenorrhyncha, phenotypic variation, behaviour, vibrational communication, troglomorphy, cave adaptation, climatic relict hypothesis, adaptive shift hypothesis, western Palaearctic

Introduction Trigonocranus emmeae was originally described by FIEBER (1876) from Switzerland. It is widely distributed in the southern half of continental Europe and Great Britain (for detailed information on its geographic distribution see HOLZINGER et al. 2003, NICKEL 2003, SELJAK 2004, MALENOVSKY & LAUTERER 2010, and MUSIK et al. 2013). Despite its comparatively large geographic range, Trigonocranus emmeae is regarded as a locally rare species; usually only a few specimens are collected, mostly from pitfall traps. Consequently, information on its ecology is sparse, but the species is assumed to “live in the top soil and leaf litter in at least moderately warm sites with incomplete vegetation cover, feeding on roots of shrubs” (NICKEL 2003: 27). In Germany, hitherto only three autochthonous populations (from Warburg, Hildesheim and Herne: NICKEL 2003) have been reported, “all from sun-exposed embankments or hillsides with scattered shrubs on damp to moderately dry soils” (NICKEL 2003: 27). Adults have been reported to display a conspicuous polymorphism concerning the size of the compound eyes, length of tegmina and bodily pigmentation (NICKEL 2003, HOLZINGER et al. 2003). A recent discovery of an apparently autochthonous Trigonocranus emmeae population in southern Germany (Rhineland-Palatinate: Mosel valley, Bernkastel) allowed us to document nymphal and varying adult phenotypes and – for the first time – a recording of their intraspecific communication signals.

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Material and methods Material examined. 6 males (all brachypterous), 18 females (16 macropterous, 2 brachypterous), 5 nymphs (1 IV. instar; 4 V. instar), Germany, Rhineland-Palatinate, Bernkastel/Mosel, from roots of Convolvulus arvensis L. (Convolvulaceae), M. Maixner leg., in coll. Museum für Naturkunde, Berlin. Subsequent to behavioural experiments, specimens were preserved dry on card mounts or in vials containing 96% ethanol. Scanning electron microscopy. For scanning electron microscopy, specimens were dehydrated with ascending ethanol series and transferred into absolute ethanol, then dried at the critical point (BAL-TEC CPD 030, BAL-TEC AG, Balzers, FL) and sputter-coated (gold/palladium, Polaron Shutter Coater SC7640); micrographs were taken with a Zeiss EVO LS10 (Carl Zeiss, Jena, DE; tungsten cathode, manufacturer’s software) at the Museum für Naturkunde, Berlin. Figures were edited with Adobe® Photoshop® CS4 and Adobe® Illustrator® CS4 (Adobe Systems Incorporated, San Jose, USA). Recording and analysis of vibrational signals. Experiments were carried out at the Museum für Naturkunde, Berlin, between June 12th–14th, 2012 (Fig. 3). Fresh roots of the host plant, Convolvulus arvensis, were placed on moist filter paper lining the floor of a 10 x 7 x 4 cm translucent plastic box with a lid. The roots were partially covered by a black plastic tube in order to imitate natural soil cavities. A small piece of reflecting foil (ca. 2 x 2 mm) was placed on the root part outside the black plastic tube. Vibrational signals emitted by Trigonocranus specimens were detected with a portable digital laser vibrometer PDV-100 (Polytec GmbH, Waldbronn, Germany) and registered directly with a Roland Edirol R-4 Pro recorder at a sampling rate of 16 bits and 44.1 kHz. Recordings were made at room temperature (22–33°C). Signal analysis was carried out using Raven Pro 1.4 (Bioacoustics Research Program, 2011). The sound files obtained are deposited at the Animal Sound Archive (TSA: Tierstimmenarchiv), Museum für Naturkunde, Berlin, Germany; accession number TSA Trigonocranus emmeae_DIG0168_02.

Results Presumed host. Nymphs and adults of Trigonocranus emmeae were found in the top soil layer of a temporarily uncultivated patch in a private garden, apparently feeding on roots (and stems close to the soil) of Convolvulus arvensis (M. Maixner, personal communication), and were collected on June 10, 2012. By this time, females had apparently already mated and were ready to oviposit as their abdomina were swollen with eggs. Phenotypic variation (Figs 1–2). Adults of Trigonocranus emmeae display remarkable intraspecific variation in regard to the overall size of the compound eyes, the number of ommatidia, length of tegmina, and bodily pigmentation. The observed variation is accentuated by sexual dimorphism, as “reductive” characters appear to be more strongly expressed in males. In the sample studied – which did not contain macropterous males – the number of ommatidia varied between 87 (in a brachypterous male) and 194 (in a macropterous female).

156 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Acoustic communication in Trigonocranus emmeae Fieber

Fig. 1. Habitus of Trigonocranus emmeae: a – brachypterous male; b – macropterous female; c – fifth instar nymph (photographs by J. Deckert, Berlin).

Fig. 2. Variation in the size of the compound eyes in Trigonocranus emmeae (SEM): a – brachypterous male; b – same, at higher magnification; c – macropterous female; d – same, at higher magnification.

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Fig. 3. Experimental set-up for recording of vibrational signals, Museum für Naturkunde: a – digital laser vibrometer and recording chamber; b – close-up of the recording chamber with root and dark zone. Arrows: reflecting foil (photographs by M. Kretzschmar, Berlin).

Phototaxis. Due to the small sample size this observation must be regarded as preliminary, however all brachypterous males tested displayed a distinct negative phototactic behaviour. Once placed inside the recording box, the brachypterous males immediately moved into the part covered by the black plastic vial (Fig. 3). On the contrary females, most of which were macropterous, were observed most frequently sitting on the underside of the translucent lid of the box, perhaps orientating towards the ceiling light. Acoustic communication. For recording experiments, one to several males were placed together with up to three females into the recording box. Within a few minutes, males were observed to initiate calling. We obtained calls from three males; females did not emit any calls, and under no constellation was active courtship leading to copulation observed. Male calls were emitted at varying intervals (less than one minute to several minutes). Single calls consisted of more or less homogenous pulse trains (Fig. 4). Total length of single calls varied from 9.88 to 27.5 s (mean: 16.7 s; SD: 5.36; n=10; N=3). The average centre frequency was 662 Hz (SD: 140; n=10; N=3). The number of pulses within a single call varies from 19 to 161 (mean: 98; SD: 32.44; n=10; N=3). The length of a single pulse is 92 ms and is very consistent (SD: 10 ms; n=20; N=3).

Discussion The large range of phenotypic variation in Trigonocranus emmeae is exceptional among Cixiidae and raises a number of questions. Reduction of compound eyes, tegmina and bodily pigmentation is usually found in species adapted to subterranean environments. Although such troglomorphies are not a priori adaptive, they are indicative of a cryptic way of life, usually associated with habitats characterized by high humidity and reduced light, such as leaf litter, crevices inside the soil, or caves. It can

158 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Acoustic communication in Trigonocranus emmeae Fieber

Fig. 4. Oscillogramm illustrating the time-amplitude pattern of a single male call of Trigonocranus emmeae (brachypterous male; in coll. Museum für Naturkunde, Tierstimmenarchiv, Berlin).

therefore be assumed that the range of morphological variation within the species is paralleled by variation of ecological parameters characterizing the species’ habitat. With nymphs living underground while adults either complete their life cycle in the top layer of the soil, or live and disperse in epigean environments, T. emmeae is – according to the classification of subterranean animals proposed by SKET (2008: 1560) – a subtroglophile (= “species inclined to perpetually or temporarily inhabit a subterranean habitat but is intimately associated with epigean habitats for some biological function (daily, e.g., feeding, seasonally, or during the life history, e.g., reproduction)”) or a eutroglophile (= “essentially epigean species able to maintain a permanent subterranean population (which may become troglobitic)”). Among the Fulgoromorpha, independent evolutionary lineages have colonized subterranean environments in many parts of the world (for a review see HOCH 2002, HOCH & WESSEL 2006, and references therein). The question as to which factors drive an epigean species to colonize underground environments has long intrigued evolutionary biologists. However, in order to study the process of cave adaptation in a natural system suitable model systems to test existing hypotheses need to be identified. The evolution of obligate cave species (troglobites) has traditionally been explained by the so-called climatic relict hypothesis (CRH) (e.g., BARR 1968). Under this hypothesis severe climatic oscillations, such as glaciation, are assumed to cause epigean populations to become extinct or extirpated, eventually resulting in the isolation of troglophilic populations in underground environments which subsequently evolve into obligate cave dwellers characterized by reduction and even loss of eyes, bodily

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 159 H. HOCH ET AL. pigmentation, and – in the case of insects – wings. This hypothesis was supported by the seeming absence of troglobites from the tropics (e.g., BARR 1968). The discovery of rich and diverse cave faunas on young oceanic islands comprising many terrestrial obligate cave-dwellers has, however, challenged this view. HOWARTH (1986) provided an alternative hypothesis on the evolution of terrestrial troglobites, which has since become known as the adaptive shift hypothesis (ASH). While the CRH essentially postulates allopatric speciation between epigean and cave-dwelling species, the ASH assumes parapatric speciation, consistent with the observed occurrence of closely related species in close geographic vicinity, e.g., on the same island. It is possible, though, that even many temperate-zone troglobites also evolved in parapatry through adaptive shifts, and that their current isolation occurred only after cave adaptation (HOWARTH & HOCH 2005, HOCH in press). HOWARTH & HOCH (2005) identified a number of intrinsic and extrinsic factors underlying adaptive shifts into cave habitats. Among the intrinsic factors are preadaptation (exaptation, i.e., ability to survive in dark, damp microhabitats), genetic repertoire (i.e., variability, adaptability to changing environments), and mating behaviour (i.e., the ability to locate and recognize potential mating partners in absolute darkness). Among extrinsic factors favouring adaptive shifts are the presence of cavernicolous habitat and exploitable food resources. The newly discovered population of Trigonocranus emmeae in Bernkastel largely fulfils the postulated requirements: the nymphs and brachypterous adults live in dark, damp soil crevices, the observed phenotypic variability suggests underlying genetic variability, and vibrational signals are a well-known communication channel in (epigean and) obligately cavernicolous Cixiidae (HOCH & HOWARTH 1993, HOCH & WESSEL 2006). Potentially suitable cavernicolous habitats are probably available, as in the Mosel valley, adjacent mountain slopes consist of loose slate (MEYER 1988), such that the existence of an extended superficial underground compartment or MSS (milieu souterrain superficiel, as described by JUBERTHIE et al. 1980) can be assumed. Under an assumed scenario of disruptive selection (either through the availability of a novel habitat or food resources, or by ecological/climatic deterioration on the surface) Trigonocranus emmeae is a promising candidate for giving rise to obligately cavernicolous species, equally conceivable in either allopatry or sympatry. The discovery of a permanent and accessible population of Trigonocranus emmeae is the first step towards establishing a new model system to study the process of cave- adaptation on the genetic, behavioural and evolutionary-developmental level in a natural system. Further research is needed to address questions of behavioural differentiation among brachypterous and macropterous individuals, symmetry of mating success among similar and different phenotypes, as well as the deciphering of the genetic and developmental mechanisms underlying phenotypic variation.

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Acknowledgements We would like to express our sincere thanks to Michael and Arntrud Maixner, Bernkastel, for collecting the specimens and for transporting them to Berlin, respectively, and for their willingness to undertake all measures for the conservation of this population in their private garden. We also thank Herbert Nickel, Göttingen, and Werner Holzinger, Graz, for information on the phenology of the species, to Moritz Kretzschmar, Berlin for the photograph of the experimental set-up (Fig. 3) and to our colleagues from the Museum für Naturkunde, Berlin: Jürgen Deckert for providing all habitus photographs, Anke Sänger for her competent and friendly assistance during scanning electron microscopy, and Manfred Asche for his constructive comments on the manuscript. Last but not least, we thank Igor Malenovský, Moravian Museum, Brno, for inviting our contribution to the present volume.

Dedication This contribution is dedicated to Pavel Lauterer on the occasion of his 80th birthday in recognition of his outstanding contribution to the taxonomy and zoogeography of the Hemiptera in Europe.

Zusammenfassung Eine kürzlich entdeckte und offenbar autochthone Population von Trigonocranus emmeae Fieber, 1876 in Deutschland (Rheinland-Pfalz: Bernkastel) bestätigte die in der Literatur erwähnte breite Spanne intraspezifischer Variabilität hinsichtlich der Größe der Komplexaugen, der Flügellänge und der Körperpigmentierung. Erstmals konnten die intraspezifischen Kommunikationssignale dieser Art aufgezeichnet werden. T. emmeae wird als mindestens subtroglophil eingeschätzt; die Eignung der Art als möglicher Modellorganismus zur Untersuchung des evolutiven Übergangs zum Höhlenleben in einem natürlichen System wird diskutiert.

References

BARR T.C. Jr. 1968: Cave ecology and the evolution of troglobites. Evolutionary Biology 2: 35–102. FIEBER F. X. 1876: Les Cicadines d’Europe d’après les originaux et les publications les plus récentes. Deuxième partie. Revue et Magasin de Zoologie (3)4: 11–268. HOCH H. (in press): Trirhacus helenae sp. n., a new cave-dwelling planthopper from Croatia (Hemiptera: Fulgoromorpha: Cixiidae). Deutsche Entomologische Zeitschrift 60(2). HOCH H. 2002: Hidden from the light of day: planthoppers in subterranean habitats (Hemiptera: Auchenorrhyncha: Fulgoromorpha). Denisia 4, Kataloge des OÖ. Landesmuseums, N.F. 176: 139–146. HOCH H. & HOWARTH F.G. 1993: Evolutionary dynamics of behavioral divergence among populations of the Hawaiian cave-dwelling planthopper Oliarus polyphemus (Homoptera: Fulgoroidea: Cixiidae). Pacific Science 47: 303–318. HOCH H. & WESSEL A. 2006: Communication by substrate-borne vibrations in cave planthoppers (Auchenorrhyncha: Hemiptera: Fulgoromorpha: Cixiidae). Pp. 187–197. In: DROSOPOULOS S. & CLARIDGE M. F. (eds.): Insect Sounds and Communication. CRC – Taylor & Francis, Boca Raton, London, New York, 532 pp.

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HOLZINGER W. E., KAMMERLANDER I. & NICKEL H. 2003: The Auchenorrhyncha of Central Europe. Die Zikaden Mitteleuropas. Vol. 1. Fulgoromorpha, Cicadomorpha, excl. Cicadellidae. Brill, Leiden, Boston, 673 pp. HOWARTH F. G. 1986: The tropical cave environment and the evolution of troglobites. Proceedings of the 9th International Congress of Speleology (Barcelona, Spain) 2: 153–155. HOWARTH F. G. & HOCH H. 2005: Adaptive shifts. Pp. 17–24. In: Culver D. C. & White W. B. (eds.): Encylopedia of Caves. Elsevier Academic Press, 654 pp. JUBERTHIE C., DELAY B. & BOUILLON M. 1980: Extension du milieu souterrain en zone non calcaire. Mémoires de Biospéologie 7: 19–52. MALENOVSKÝ I. & LAUTERER P. 2010: Additions to the fauna of planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) from the Czech Republic. Acta Musei Moraviae, Scientiae biologicae 95(1): 49–122. MEYER W. 1988: Geologie der Eifel. 2. ergänzte Aufl. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, 615 pp. MUSIK K., WALCZAK M., DEPA £., JUNKIERT £., JEDYNOWICZ A. 2013: Trigonocranus emmeae Fieber, 1876 (Hemiptera, Fulgoromorpha, Cixiidae) – a new species for Poland. In: POPOV A., GROZEVA S., SIMOV N., TASHEVA E. (eds.): Advances in Hemipterology. ZooKeys 319: 249–253. NICKEL H. 2003: The Leafhoppers and Planthoppers of Germany (Hemiptera, Auchenorrhyncha): Patterns and strategies in a highly diverse group of phytophagous insects. Pensoft Publishers, Sofia, Moscow, and Goecke & Evers, Keltern, 460 pp. SELJAK G. 2004: Contribution to the knowledge of planthoppers and leafhoppers of Slovenia (Hemiptera: Auchenorrhyncha). Acta Entomologica Slovenica 12: 189–216. SKET B. 2008: Can we agree on an ecological classification of subterranean animals? Journal of Natural History 42(21): 1549–1563.

162 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 163–174, 2013

The South-East Asian planthopper genus Lanaphora, with description of a new species (Hemiptera: Fulgoroidea: Delphacidae: Tropidocephalini)

MANFRED ASCHE, Fls Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Hemiptera Research Group, Invalidenstraße 43, 10115 Berlin, Germany and Research Associate, Department of Entomology, Bishop Museum, P.O. Box 19000-A, Honolulu, Hawai’i 96817, U.S.A; e-mail: [email protected]

ASCHE M. 2013: The South-East Asian planthopper genus Lanaphora, with description of a new species (Hemiptera: Fulgoroidea: Delphacidae: Tropidocephalini). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 163–174. – The hitherto monotypic South-East Asian planthopper genus Lanaphora Muir, 1915 is revised. Lanaphora bakeri Muir, 1915 from the Philippine Islands is redescribed and a new species from Papua New Guinea, Lanaphora paveli sp.nov., is described; both species are illustrated including details of the male genitalia. A third species is reported from the Solomon Islands but not formally described here. All Lanaphora species are bamboo-feeders and may be potential pests. Keywords. Auchenorrhyncha, Fulgoromorpha, , taxonomy, excessive female wax production, bamboo-feeder, Philippine Islands, Papua New Guinea, Solomon Islands

Introduction The Delphacidae of South East Asia must still be regarded as insufficiently known. Despite recent monographies from Taiwan (YANG & YANG 1986, YANG 1989) and China (DING 2006), vast areas of this immense land and island realm remain virtually untouched by contemporary faunistic and taxonomic studies. Concerning the Tropidocephalini of this area only a few taxonomists have contributed, mostly in the early 20th century. Tropidocephaline taxa were predominantly reported from the Philippine Islands, e.g. by MUIR (1913, 1915, 1916, 1919), from Malaysia, e.g. by MATSUMURA (1907) and MELICHAR (1914), from Indonesia by e.g. by BIERMAN (1908), MUIR (1913) and SCHMIDT (1926), and – more “recently” – from Indonesia: Sunda Archipelago by JACOBI (1941) and New Guinea by FENNAH (1965). Among all South East Asian Tropidocephalini, the genus Lanaphora Muir, 1915 is one of the least known. Since its monotypic establishment with Lanaphora bakeri Muir, 1915 from the Philippine Islands: Luzon, based on one male and one female (MUIR 1915), it was subsequently listed by MUIR (1916) without providing new data, and mentioned again by MUIR (1917) in a comparison with Vizcaya Muir, 1917 (today in the subfamily Vizcayinae, see ASCHE 1990) stating superficial similarity. For further listing see METCALF (1943). Lanaphora bakeri has never been illustrated. In the course of studies on Oriental Delphacidae I was able to examine the type material of L. bakeri in the collections of the Bishop Museum, Honolulu, and also discovered additional conspecific specimens from the Philippines. L.

163 M. ASCHE bakeri is re-described and for the first time genitalia are illustrated here; a lectotype is designated. The collections of Bishop Museum and the Natural History Museum, London, also contained Lanaphora specimens from Papua New Guinea and the Solomon Islands which could not be assigned to L. bakeri, and apparently represent hitherto undescribed species. The species from Papua New Guinea is described below as Lanaphora paveli sp.nov., the specimen from the Solomon Islands, a female, is doubtlessly a congener; however, due to its poor condition it is not described here as a new species. The relationships of Lanaphora to other Tropidocephalini are briefly discussed.

Material and methods Measurements and illustrations (line drawings and photographs) of external body parts were taken from dry specimens. The terminology of bodily structures, including male and female genitalia, follows ASCHE (1985). A Leitz stereomicroscope with camera lucida attachment was used for morphological examinations and for producing line drawings. For studying the male genitalia the abdomina were removed from the specimens and macerated in 10% KOH for 24 hours at room temperature, then transferred to glycerin for storage and/or to glycerin-jelly for preparing drawings. Photographs resulted from combined image stacks obtained by a camera (Canon EOS 450 D) attached to a Leica MZ 16 stereo microscope and Combine ZP software; subsequently they were processed with Adobe Photoshop CS 3.

Specimens examined are deposited in the following collections: BMNH ...... The Natural History Museum, London, UK BPBM ...... Bernice P. Bishop Museum, Honolulu, Hawai´i, USA

Taxonomy Lanaphora Muir, 1915 Lanaphora Muir, 1915: 317. Type species: Lanaphora bakeri Muir, 1915, by original designation and monotypy. Supplementary description. Small-sized tropidocephaline delphacids with delicate appearance; tegmina steeply tectiform, antennal joints (especially scape) elongate and slightly compressed, lateral margins of frons foliately ridged. Body colouration generally whitish or light yellow to brownish; head, antennae and tegmina with distinct brown or orange spots or markings. Head including compound eyes distinctly narrower than pronotum; vertex subtriangular, strongly narrowing towards apex, lateral carinae foliately ridged, median carina weak, surface of vertex deeply concave, transition of vertex to frons limited by a ridged transverse carina. Frons elongate and narrow, slightly widening towards frontoclypeal suture; lateral carina foliately ridged; median carina distinctly present in lower two thirds, then vanishing towards apex. Postclypeus short with ridged

164 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Lanaphora Muir lateral and median carina, anteclypeus devoid of carinae. Rostrum attaining posterior margin of middle coxae. Antennal joints subequal in shape, scape elongate, slightly compressed, pedicel terete and distinctly shorter than scape; pedicel with numerous sensory plaques. Pro- and mesonotal carinae straight, attaining posterior margin, lateral carinae slightly diverging; pronotal carinae strongly ridged. Tegmina elongate, widest distally of nodal line, surpassing the tip of abdomen by about half their length; apical margin slightly undulate and tapering. Hind wings distinctly shorter than tegmina, like in tropidocephaline delphacids with anastomosis of M and Cu. Post-tibial spur slender, posterior margin smooth, devoid of teeth, two thirds as long as postbasitarsus, inner margin with very fine setae; postbasitarsus distally with 6–7 teeth in a slightly angulate row, second posttarsal joint distally with 4 teeth in a closed row. Male genitalia with genital segment laterocaudally broadly or bulbously rounded; ventrocaudal margin of genital segment medially with a bifid process. Aedeagal complex basally embraced by a relatively short, hood-shaped anal segment (lacking processes); aedeagus in basal third or at midlength subrectangularly bent ventrally, with a single spinose process on the right side. Parameres slender, elongate, erect dorsally, in repose parallel to diaphragm, attaining or surpassing anal segment. Female genitalia with relatively short and sturdy ovipositor which does not attain the tip of abdomen; lateral gonapophyses IX strongly compressed and dilated, with excessive wax-production. Differential diagnosis. Lanaphora can readily be distinguished from other tropidocephaline genera which display undulate and pointed tips of the tegmina like Arcofacies Muir, 1915 and Arcofaciella Fennah, 1956 by the foliately ridged lateral frontal carinae (versus not foliate), by the shape and proportions of the antennal joints (scape longer than pedicel and compressed versus shorter and terete). Lanaphora differs from Purohita Distant, 1906 especially by the shape of the scape (elongate and slightly compressed in Lanaphora, strongly dilated and foliately compressed in Purohita). From the Chinese genus Neocarinodelphax Chen & Tsai, 2009 which also displays a frons with strongly ridged lateral carinae, it differs by the shape of the antennae (scape and pedicel shorter and terete in Neocarinodelphax) and by the shape of the genital segment (laterally on each side with a strongly projected process in Neocarinodelphax versus no such process in Lanaphora). Distribution. Only known from South East Asia: Philippine Islands, New Guinea, and the Solomon Islands: Bougainville. A wider distribution in this region is assumed. Remarks. Within the Tropidocephalini, Lanaphora remains isolated. In the key for the Chinese Tropidocephalini provided by QIN & ZHANG (2010) Lanaphora would belong to a small group with a subtriangular vertex like Conocraera Muir, 1916 and Tropidocephala Stål, 1853; however, the unique bodily features including those of the male and female genitalia do currently not suggest a closer relationship to any of these or other genera. Muir’s generic name “Lanaphora” may be a bit misleading as the Latin “lana” means wool. It is very likely that Muir referred to the woolly appearance of excessive wax-production of the females.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 165 M. ASCHE

Lanaphora bakeri Muir, 1915 (Figs 1–5, 11–14) Lanaphora bakeri Muir, 1915: 317.

Type material. Lectotype ♂ (macropterous), here designated, Philippine Islands: Luzon, pinned together with a ♀ on the same block, with labels [white] “Maquilin, Luzon” [handwritten], “Lanaphora bakeri Muir” [handwritten by Muir], “930”, [red] “Type of [printed] Lanaphora Muir [handwritten by Muir]”, “Type of [printed] L. bakeri Muir [handwritten by Muir]”, “lectotype = ♂, det. Asche, 1984” (deposited in BPBM, # 4855). Paralectotypes: 1 ♀, on the same mount as lectotype, “paralectotype = ♀, det. Asche, 1984”; 1 ♀, same data as lectotype. Both in BPBM. Additional material examined. 1 ♂, Philippine Islands: Luzon, Camarines Sur, Mt. Isarog, Pili, 800–900 m,12.iv.1965; 1 ♀, ibid., 500 m, 4.iv.1963; both: H. M. Torrevillas leg. 2 ♂♂, 1 ♀, ibid., Los Baños, Baker leg. 1 ♀, Negros, Negros Or., Mt. Talinas, 900 m, in rainforest, 29.vi.1958, H. E. Milliron leg.. All in BPBM. Supplementary description. The colouration of body and tegmina has been excellently documented by MUIR (1915: 317–318). For structure and proportions of head and thorax including antennae, tegmina, wings and legs see the generic diagnosis. Male genitalia. Genital segment in lateral aspect trapezoidal, ventrally about 2.4 times longer than dorsally; laterodorsal margin evenly convex, lateroventral margin on each side projected into a distinct bulbus; medioventral margin with a slender, apically bifid process; lateral margins rounded towards diaphragm, the latter covering more than two thirds of the caudal surface; dorsal margin of diaphragm concave, foramen for parameres broadly oval with a small triangular median projection at the ventral margin; laterocaudal margin of genital segment and dorsolateral part of diaphragm with distinct setae. Anal segment relatively short, about 1.5 times longer than wide, subcylindrical with ventral surface membraneous and concave, ventrocaudally on each side slightly projected, ventrobasally on each side arm-like expanded and ventrally embracing the aedeagal base; anal segment furnished with numerous strong setae, especially at ventrocaudal margin; anal style slightly shorter than anal segment. Shaft of aedeagus slightly twisted at base, at about mid-length subrectangularly bent ventrally; basal part strongly sclerotized and voluminous, on inner ventral side a sturdy, concave socket; distal part slender, in lateral aspect slightly sinuate, phallotreme subapically on caudal side; in right lateral aspect a slender, terete spine dorsally emerging from a deep furrow and curved ventrally to the right side of aedeagus, not surpassing half the length of the bent distal part. Parameres elongate, distally attaining base of anal segment; in ventrocaudal aspect inner margin straight, outer margin slightly convex, distally produced into a short lobe-like projection which is directed laterocaudally. Female genitalia. Ovipositor short and sturdy, not reaching the base of the anal segment; gonapophyses IX strongly dilated, dorsoventrally flattened at base, at about midlength narrowing and sides bent dorsally causing a distinct edge, distal sides of gonapophyses IX laterally compressed with excessive wax production. Valvifers VIII slender, widening at base, inner basal margins touching each other. Anal segment very short, about 3 times wider than long, slightly depressed; anal style little longer than anal segment, lanceolate with ventral surface slightly concave. Measurements. Body length: males (n=3): 3.8–4.4 mm, females (n=3): 4.7–5.0 mm. Differential diagnosis. Lanaphora bakeri is similar in most characters to L. paveli sp.nov. from New Guinea described below but differs considerably in the following

166 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Lanaphora Muir

Figs 1–5. Lanaphora bakeri Muir. 1 – male genitalia, left lateral aspect; 2 – male genitalia, ventrocaudal aspect; 3 – male genital segment, ventral aspect; 4 – male anal segment, aedeagus complex and parameres, right lateral aspect; 5 – parameres, ventrocaudal aspect. Scale bar 0.1 mm.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 167 M. ASCHE details: Colouration of L. paveli predominantly orange to light brown, scape anteriorly nearly entirely orange. Male genital segment ventrally with distinct laterocaudal bulbous projections, medioventral process rather slender and only terminally bifid. Parameres attaining lower margin of the anal segment. Aedeagus at level of bending ventrally with a sturdy concave socket, aedeagal spine not surpassing mid-length of distal part. Distribution. Philippine Islands: Luzon (MUIR 1915, 1916), Negros (new record). Host plant and ecology. In the original description MUIR (1915: 318) reports that the specimens he examined were “living at base of leaf-sheath of bamboos among a white, flocculent secretion”. According to label information of additional specimens examined here they were collected in elevations between 500 and 900 m, the female from Negros in a rainforest. Remarks. This species was named by Muir in honour of the American botanist and entomologist Charles Fuller Baker (1872–1927) who that time was employed at the College of Agriculture of the University of the Philippines in Los Baños, Luzon. Baker himself worked mainly on Cicadelloidea, but in his later years also on Fulgoroidea. Baker did extensive field work in the Philippines and other parts of South East Asia, and generously provided colleague specialists like Frederick Muir with valuable specimens. Baker had observed and collected the first Lanaphora specimens in the Mount Maquiling area and shared this information with Muir (see MUIR 1915: 318).

Lanaphora paveli sp.nov. (Figs 6–10, 15–17)

Type material. Holotype ♂ (macropterous), Papua New Guinea, Simbu P. Karimui, 1100 m, on young shoots of Bambusa sp., heavy infestation, 10.v.1984, I.W. Ismay leg. (deposited in BMNH). Paratypes: 1 ♂, 3 ♀♀, same data as holotype (BMNH); 2 ♂♂, 3 ♀♀, Papua New Guinea, NE, Tapo (= Tapu), 3 km NW of Kainatu, 1650 m, 22.x.1959, T. C. Maa leg. (BPBM). Description. Coloration. Very lightly coloured species, yellow to whitish, with orange to brown sprinkles or marks, especially on head, antennae, tegmina and legs. Vertex including carinae white except for a narrow blackish brown infuscation at the posterior ridges of lateral carinae caused by a dark brown stripe on the sides above compound eyes. Frons white to pale yellow, foliate lateral carinae white with 4 narrow brown stripes on outer sides. Post- and anteclypeus pale yellow. Antennal joints with ground color white; scape anteriorly from upper base to lower middle a narrow oblique dark brown to blackish stripe, distal tip anteriorly orange-brown; pedicel in proximal half white with a narrow oblique dark brown ring, in distal half sordid brown. Rostrum with a brown tip. Genae and head laterally anteriorly and dorsally of compound eyes white. Pronotum white with orange patches behind compound eyes and between lateral and median carinae. Mesonotum orange-white with brown marks at the anterior and posterior end of the lateral carinae; tip of scutellum with a narrow transverse brown band. Tegulae white. Tegmina translucent, slightly opaque; veins and pustules on veins concolorous, between the pustules numerous more or less extended brown spots; in about mid-length of anterior part before nodal line between ScR and Cu1 an extended brown-orange marking which forms a nearly closed circle towards Cu1; at level of nodal line from the outer discoidal

168 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Lanaphora Muir cell towards inner margin an oblique brown-orange stripe; in distal part between M1 and M2 a longitudinal brown mark; all marginal veins from Sc1 towards Cu1a with a brown fringe. Hind wings translucent, veins sordid brown. Legs white to pale yellow, tibiae with 2–3 narrow ring-like brown marks, fore- and middle tarsi brown. Abdominal tergites brown, sternites in males pale brown, in females white. Male genitalia with genital segment yellowish except for a black medioventral process; anal segment and anal style pale yellow; parameres and aedeagus melichrous. Female genitalia with ovipositor and anal segment pale yellow, anal style whitish. Structure. Head including compound eyes 0.6 times narrower than pronotum. Vertex narrow, subtriangular, posterior margin 1.25 times wider than long in midline and 2.15 times wider than anterior margin; lateral margins strongly ridged, foliate; median carina and limited compartments absent, area of vertex deeply concave, limitation towards frons v-shaped with a ridged transverse carina. Frons slender, very narrow at apex, widening towards frontoclypeal suture, medially about 2.6 times longer than wide at base and about 1.25 times longer than post- and anteclypeus together; lateral frontal carinae strongly foliately ridged, over most of their length slightly convex, almost straight, only in lower part close to frontoclypeal suture distinctly diverging; median frontal carina ridged and weakly forked at apex; frontal surface shallowly concave. Postclypeus vaulted, lateral and median carinae sharply ridged; anteclypeus rounded, devoid of lateral carinae, median carina faintly visible in upper part. Antennal joints elongate, almost equally long; scape compressed, anteriorly and posteriorly slightly concave with upper and lower margin roundly ridged, distally expanding: about twice as wide than at base; pedicel terete, distal half covered with about 25 sensory plaques, irregularly arranged. Genae convex, oblique carina ridged, vanishing towards antennal base. Compound eyes flat kidney-shaped with a deep incision for the upper antennal base; ocelli relatively small. Pronotum about as wide as mesonotum; carinae strongly ridged; lateral carinae straight, diverging, attaining posterior margin; surface of disk concave. Mesonotum in midline about 5 times longer than pronotum, disk only slightly vaulted, almost plain; lateral carinae nearly straight, diverging, at posterior end slightly curved laterally, vanishing in a swelling and not reaching posterior margin; median carina mostly ridged, fading at anterior end of scutellum, then strongly ridged towards posterior tip. Tegulae well developed. Tegmina elongate, narrow at base, widening towards apex, about 3 times as long as maximally wide, widest at level of Cu1a; nodal line in about mid-length of tegmen; inner discoidal cell distinctly larger than outer one; hind margin slightly undulate with a pointed tip at level of M1; inner margin of clavus at level of entry of common branch of claval veins foliately ridged; veins slightly prominent, beset with numerous very fine callous pustules, however devoid of setae or bristles. Legs slender, fore- and middle femora and tibiae slightly compressed; post-tibiae about 1.4 times longer than post-tarsal joints together, subquadrangular in cross-section with strongly developed longitudinal rims, enclosed surface slightly concave; laterally with 2 small spines, distally with 3 sturdy outer and 2 smaller inner teeth; post-basitarsus distally with 6 teeth in an even row, 2nd post-tarsal joint distally with 4 small teeth in a row; post-tibial spur about 0.7 times shorter than post-basitarsus, surface vaulted but slightly flattened, devoid of marginal teeth, numerous fine setae at inner margin.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 169 M. ASCHE

Male genitalia. Genital segment trapezoidal with laterodorsal margin broadly rounded, ventrally about 3.3 times longer than dorsally, lateroventrally on each side a small subtriangular but rounded projection, ventrocaudal margin medially furnished with a broad and sturdy deeply bifurcate process; diaphragm covering about two thirds of the caudal surface, dorsal margin deeply emarginate; foramen for parameres flat ovate; laterocaudal margins of pygofer opening smoothly rounded medially and beset with sturdy bristles, diaphragm only little sunk cephally. Anal segment relatively short, tubular with ventral surface concave and membranous, about 1.4 times longer than wide, ventrocaudally slightly projected, rounded, ventrobasally projected into membranous arms embracing the aedeagal base; anal style lanceolate, about 0.4 times smaller than anal segment. Shaft of aedeagus at one third of its length measured from base abruptly bent ventrally in an almost right angle; on the ventral right side of bend, a small semi- shell-like projection; dorsally on the right side a slender curved spine emerging from a deep furrow which nearly parallels direction of bent aedeagus and reaches to half-length of its distal part; the latter on the right side nearly over the whole length with a narrow, slightly sinuate rim; phallotreme subapically on the caudal side. Parameres slender and extremely elongate, attaining (in repose even surpassing) the ventrocaudal margin of the anal segment, in ventral aspect slightly sinuate, broad at base with a small hook-like process, distally narrowing and converging, apically diverging and dilated with truncate tip pointing laterally. Female genitalia very similar to those of L. bakeri. Valvifers VIII relatively broad, dilated at base, base medially with lip-like projection. Measurements. Body length: males (n= 4): 4.4–4.6 mm, females (n=6): 4.9–5.1 mm. Differential diagnosis. Lanaphora paveli sp.nov. can be distinguished from L. bakeri by the following external and genitalic characters: scape of antenna anteriorly whitish with a narrow oblique dark brown stripe (versus yellowish with a broad orange-brown band in L. bakeri), tegmina in anterior half with an extended dark brown spot almost forming a circle (versus only brown patches without such circle in L. bakeri), male genital segment ventrolaterally with a small subtriangular projection (versus a distinct bulbous projection in L. bakeri), ventrocaudal margin of genital segment medially with a very broad and deeply bifurcate process (versus a narrow and only apically bifid process in L. paveli), anal segment ventrocaudally broadly rounded (versus with small projections on each side in L. paveli), aedeagus ventrally on the right side at level ofbending with a small husk- like projection (versus a sturdy socket in L. paveli), aedeagal spine reaching mid-length of distal part (versus not reaching mid-length in L. paveli), parameres in repose attaining or surpassing ventrocaudal margin of anal segment (versus not attaining or surpassing this margin in L. bakeri), in ventrocaudal aspect median margin of parameres distinctly concave (versus straight in L. paveli). Etymology. This species is named in honour of the Czech entomologist Pavel Lauterer as an expression of sincere respect for his life-time achievements in taxonomy and faunistics of Auchenorrhyncha as well as of Psylloidea. Distribution. Papua New Guinea.

170 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Lanaphora Muir

Figs 6–10. Lanaphora paveli sp.nov. 6 – male genitalia, left lateral aspect; 7 – male genitalia, ventrocaudal aspect; 8 – male genital segment, ventral aspect; 9 – anal segment, aedeagus complex and parameres, right lateral aspect; 10 – parameres, ventrocaudal aspect. Scale bar 0.1 mm.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 171 M. ASCHE

Hostplants and ecology. Largely unknown. According to label information collected on Bambusa sp. (Bambusaceae). Remarks. The wide overlap in bodily configuration and apparently host plant association (bamboo-feeders) suggests that L. paveli and L. bakeri are closely related. The geographic range of each of these species, L. bakeri occurring in the Philippine islands of Luzon and Negros, L. paveli sp.nov. in Papua New Guinea, shows a considerable allopatric distribution. Despite of intensive fieldwork in many regions of South East Asia carried out from the 1960s to date by many entomologists, e.g. of the Bishop Museum Honolulu, no further specimens have become available. This is even more surprising as apparently Lanaphora species seem to be locally quite abundant (“heavy infestation” on Bamboo-shoots in New Guinea, according to label information). As Lanaphora species appear to cause damage to bamboo, their potential pest status should be assessed. It is expected that in the course of future field-work with focus on bamboo-feeders the geographic range of these two species will expand, and possibly even more Lanaphora species will be discovered.

Lanaphora sp.

Material examined. 1 ♀, Solomon Islands: Bougainville (S.), Fokure, 680 m, 13.vi.1956, E. J. Ford Jr. leg. (BPBM). Remarks. A single female specimen from the Solomon Islands: Bougainville was found in the collections of the Bishop Museum, Honolulu which is congeneric with L. bakeri and L. paveli sp.nov.; however, this individual is partly damaged and therefore not illustrated here. It differs from L. paveli sp.nov. by displaying an extended and continuous blackish stripe on the anterior side of the dilated antennal scape (oblique, short and narrow in L. paveli). Body length: 4.9 mm. It is very likely a new species; however, I shall refrain from describing it until more material, preferably including males, becomes available.

Acknowledgements In the first place I wish to thank Igor Malenovsky from the Moravian Museum in Brno for inviting my contribution to this festschrift for Pavel Lauterer and Jaroslav Stehlík. The late Wayne Gagné, Bishop Museum, Honolulu, who made the type material of Lanaphora bakeri accessible already in the mid 1980s when I was still working on my PhD thesis on the phylogeny of Delphacidae deserves my sincere thanks and memory. I also thank Gordon M. Nishida, formerly Bishop Museum, Honolulu, Michael D. Webb, the Natural History Museum, London, and Michael R. Wilson, National Museum and Galleries of Wales, Cardiff, for arranging loans of further Lanaphora specimens. I am very grateful to Manfred Uhlig, Museum für Naturkunde, Berlin for providing and processing the color images, and to Charles R. Bartlett, University of Delaware, Newark for his constructive comments on the manuscript. Thanks also to my dear wife Hannelore Hoch who accompanied this paper all along its way.

172 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Lanaphora Muir

Figs 11–17. 11–14 – Lanaphora bakeri Muir. 11 – habitus, left lateral aspect; 12 – frontal aspect; 13 – ovipositor, left lateral aspect; 14 – ovipositor with excessive wax secretion at the gonapophyses IX, left lateral aspect. 15–17 – Lanaphora paveli sp.nov. 15 – habitus, dorsal aspect; 16 – frontal aspect; 17 – habitus, left lateral aspect.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 173 M. ASCHE

References

ASCHE M. 1985: Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromorpha) Teil 1: Text, Teil 2: Abbildungen. Marburger Entomologische Publikationen 2(1, 1 & 2): 1–912. ASCHE M. 1990: Vizcayinae, a new subfamily of Delphacidae with revision of Vizcaya Muir (Homoptera: Fulgoroidea) – a significant phylogenetic link. Bishop Museum Occasional Papers 30: 154–187. BIERMAN C. J. H. 1908: Homopteren aus Semarang (Java) gesammelt von Herrn Edw. Jacobson. Notes from the Leyden Museum 29: 151–169. DING J. 2006: Homoptera Delphacidae. Fauna Sinica. Insecta Vol. 45. Science Press, Beijing, 776 pp. + 20 pls. FENNAH R. G. 1965: Delphacidae from Australia and New Zealand (Homoptera: Fulgoroidea). Bulletin of the British Museum (Natural History), Entomology 17(1): 1–59. MATSUMURA S. 1907: Monographie der Homopteren-Gattung Tropidocephala Stål. Annales Historico- Naturales Musei Nationalis Hungarici 5: 56–66. JACOBI A. 1941: Die Zikadenfauna der Kleinen Sundainseln. Nach der Expeditionsausbeute von B. Rensch. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Thiere (Jena) 74: 277–322. MELICHAR L. 1914: Neue Fulgoriden von den Philippinen: I. Theil. Philippine Journal of Sciences 9: 269–283. METCALF Z. P. 1943: General Catalogue of the Hemiptera. Fulgoroidea, Part 3. Fascicle IV, Araeopidae (Delphacidae). Smith College, Northhampton, Massachussetts, 552 pp. MUIR F. 1913: On some new Fulgoroidea. Proceedings of the Hawaiian Entomological Society 2: 237–269. MUIR F. 1915: A contribution towards the taxonomy of the Delphacidae. Canadian Entomologist 47: 208–212; 261–270; 296–302; 317–320. MUIR F. 1916: Additions to the known Philippine Delphacidae (Hemiptera). Philippine Journal of Sciences 11: 369–385. MUIR F. 1917: A new Philippine genus of Delphacidae. Philippine Journal of Sciences 12: 351–352. MUIR F. 1919: Some Malayan Delphacidae. Philippine Journal of Sciences 15: 521–531. QIN D. Z. & ZHANG Y.-L. 2010: A key to the genera of Tropidocephalini (Hemiptera: Fulgoromorpha: Delphacidae) of China with description of Mucillnata rava, new genus and species. Zootaxa 2448: 61–68. SCHMIDT E. 1926: Fauna Buruana. Homoptera. Treubia 7: 217–258. YANG CH.-T. 1989: Delphacidae of Taiwan (II) (Homoptera: Fulgoroidea). NSC Special Publications (Taipei, Taiwan) 6: 1–334. YANG J.-T. & YANG CH.-T. 1986: Delphacidae of Taiwan (I) Asiracinae and the Tribe Tropidocephalini (Homoptera: Fulgoroidea). In: Collected Papers on Homoptera of Taiwan. Taiwan Museum Special Publications 6: 1–79.

174 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 175–182, 2013

Notes on the genus Sarima (Hemiptera: Fulgoroidea: Issidae) with description of a new genus from Sri Lanka

VLADIMIR M. GNEZDILOV Zoological Institute, Russian Academy of Sciences, Universitetskaya nab.1, 199034 Saint Petersburg, Russia; e-mail: [email protected], [email protected]

GNEZDILOV V. M. 2013: Notes on the genus Sarima (Hemiptera: Fulgoroidea: Issidae) with description of a new genus from Sri Lanka. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 175–182. – A redescription of the genus Sarima Melichar, 1903 is provided. As far as is known to the author, the genus Sarima sensu stricto comprises only three species (Sarima illibata Melichar, 1903, S. elongata Melichar, 1903, and S. cretata Distant, 1906) and is limited in its distribution to Sri Lanka. A new genus, Pavelauterum gen. nov., is erected for Hysteropterum fusculum Melichar, 1903 from Sri Lanka, which was treated earlier as a member of the genus Sarima. Lectotypes are designated for Sarima illibata, S. elongata and Hysteropterum fusculum. Keywords. Auchenorrhyncha, Fulgoromorpha, Issini, taxonomy, new combination, lectotype designations, Oriental Region

Introduction The genus Sarima was erected by MELICHAR (1903) for two species, Sarima illibata Melichar, 1903 (type species) and Sarima elongata Melichar, 1903 from Sri Lanka. Later 27 more Oriental species were added to the genus (METCALF 1958; HORI 1970, 1971). The revision of the genus Sarima auctorum was started by CHAN & YANG (1994) who, describing Taiwanese fauna of the family Issidae, transferred S. rubricans Matsumura, 1916 and S. matsumurai Esaki, 1931 to the genus Eusarima Yang, 1994 (in CHAN & YANG 1994) and S. pallizona Matsumura, 1938 to the genus Parasarima Yang, 1994 (in CHAN & YANG 1994). In the present study I give a diagnosis of the genus Sarima in a restricted sense based on the characters of the type species and erect a new genus for Hysteropterum fusculum Melichar, 1903 described from Sri Lanka and later transferred by MELICHAR (1906) to the genus Sarima. Examination of type specimens of H. fusculum deposited in the Museum für Naturkunde (Berlin, Germany) shows that this species does not belong to the genus Sarima s. str. and cannot be attributed to any other known issid genus. This leads me to erect a new genus to accommodate this species. Apparently the genus Sarima s. str. is limited in its distribution to Sri Lanka. Further studies are needed to clarify how many genera are now covered under the Sarima auctorum – currently 29 species known from the Eastern Palaearctic, Oriental, and Australasian Regions are included (BOURGOIN 2012; HORI 1970, 1971). I am happy to take this opportunity to dedicate the new genus described below to Dr Pavel Lauterer, who oversaw Melichar’s collection in the Moravian Museum and studied

175 V. M. GNEZDILOV

Central European Hemiptera intensively for many years, continuing the long tradition of natural history studies in Brno.

Material and methods The morphological terminology herein follows ANUFRIEV & EMELJANOV (1988) for the head, EMELJANOV (2001) for the pronotum, and EMELJANOV (1971) for the hypocostal plate of the forewing. The drawings were made using a Leica M165C compound light microscope. The photographs were taken using a Leica Z16 APOA microscope with a Leica DFC490 video camera and produced with the Leica Application Suite ver. 3.7, Auto-Montage Essentials, and Adobe Photoshop software. The material examined is deposited in the following collections:

BMNH ...... Natural History Museum, London, United Kingdom MMBC ...... Moravian Museum, Brno, Czech Republic ZMHB ...... Museum für Naturkunde, Berlin, Germany

Taxonomy

Family Issidae Spinola, 1839 Subfamily Issinae Spinola, 1839 Tribe Issini Spinola, 1839 Genus Sarima Melichar, 1903 Sarima Melichar, 1903: 78. Type species: Sarima illibata Melichar, 1903 (by original designation). Redescription. Metope wide, slightly convex, enlarged above clypeus, with median carina running from its upper margin to metopoclypeal suture; sublateral carinae distinct only in upper half of metope (Figs 1, 8, 10). Median and sublateral carinae joint below upper margin of metope which is slightly concave or almost straight. Lateral margins of metope keel-shaped. Postclypeus slightly flattened dorso-ventrally, with no carina. Pedicel nearly spherical. Coryphe transverse, sometimes with weak median carina; anterior margin weakly convex; posterior margin obtusely angulate (Figs 9, 11). Ocelli present. Pronotum sometimes with weak median carina, anterior margin right-angled, keel-shaped and elevated, posterior margin almost straight. Paradiscal fields of pronotum very narrow behind the eyes. Paranotal lobes of pronotum wide and rounded, flat, with no carina. Mesonotum slightly longer than pronotum, with weak median and lateral carinae. Forewings elongate (Figs 3, 5, 6), with hypocostal plate. Basal cell narrowly oval, precostal area with transverse veins in distal half of wing. Radius bifurcate, dividing near to the basal cell, anterior branch (R1) short and fusing with posterior branch slightly basad wing mid-point, forming a loop (Fig. 3), median with 3 branches (dividing in distal half of wing), cubitus anterior bifurcate (dividing near wing mid-point); secondary transverse veins present in some cells (Fig. 3). Clavus as long as nearly 4/5 of wing

176 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Notes on Sarima with description of new genus

Figs 1–2. Head in frontal view: 1 – Sarima illibata Melichar, male lectotype; 2 – Pavelauterum fusculum (Melichar), female lectotype.

length, open (cubitus posterior and postcubitus + first anal vein joint apically); postcubitus and first anal vein joint at mid-point of clavus. Hind wings three-lobed, nearly as long as forewings. Hind tibia with two lateral spines in its distal half and with 6–7 apical spines. First metatarsomere with two latero-apical and 7 intermediate apical spines in complete, arc-shaped row. Female sternum VII with hind margin weakly convex. Gonoplacs convex, rounded. Female anal tube long and narrow, rounded apically, anal column short.

Sarima illibata Melichar, 1903 (Figs 1, 3, 5, 8, 9) Sarima illibata Melichar, 1903: 79.

Type material examined (MMBC). SRI LANKA: 1 ♂ (lectotype, here designated), “Ceylon, Peradeniya, 24.II.1902, leg. Dr. Uzel / Dr. Melichar (on lateral margin)” (printed, with date handwritten in ink), “Collectio Dr. L. Melichar, Moravské museum Brno” (printed), “Transcriptio (printed), Sarima illibata sp .n. ♂ (handwritten in ink), L. Melichar det. 1903 (printed, with date handwritten in ink), “Syntypus” (red, printed), “Invent. è. 3834 / Ent., Mor. muzeum, Brno” (printed, with number handwritten in ink); 1 ♂ (paralectotype), “Ceylon, Peradeniya, 9.I.1902, leg. Dr. Uzel / Dr. Melichar (on lateral margin)” (printed, with date handwritten in ink), “Collectio Dr. L. Melichar, Moravské museum Brno” (printed), “illibata (handwritten in ink) det. Melichar. (printed)”, “Typus” (red, printed), “Transcriptio (printed), Sarima illibata sp .n. ♂ (handwritten in ink), L. Melichar det. 1903 (printed, with date handwritten in ink), “Syntypus” (red, printed), “Invent. è. 3830 / Ent., Mor. muzeum, Brno” (printed, with number handwritten in ink); 1 ♀ (paralectotype), “Ceylon, Peradeniya, Jan. 1902, leg. Dr. Uzel / Dr. Melichar (on lateral margin)”, “Collectio Dr. L. Melichar, Moravské museum Brno” (printed), “illibata (handwritten in ink) det. Melichar. (printed)”, “Syntypus” (red, printed), “Invent. è. 3828 / Ent., Mor. muzeum, Brno” (printed, with number handwritten in ink).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 177 V. M. GNEZDILOV

Supplementary description. Pro- and mesonotum with median carina. Forewings with quite wide hypocostal plate. General coloration light yellow-greenish. Metope with two light brown (in males) or brown (in female) bands. Forewings with light brown corium below radius and the whole clavus light brown. Hind wings matt, with brown or dark brown veins. Leg spines black. Total length: males – 5.0 mm; females – 5.5 mm. Note. MELICHAR (1903) mentioned several localities for the type series. I have examined the specimens from Peradeniya and Heneratgoda (both in central Sri Lanka, Kandy District) collected by Dr. Uzel and deposited in MMBC. The specimen from Heneratgoda differs from the specimens from Peradeniya in coloration and may represent another species. Here I designate the lectotype for the male from Peradeniya according to the ICZN (1999: Art. 74) to stabilize the nomenclature in the genus.

Sarima cretata Distant, 1906 Sarima cretata Distant, 1906: 344.

Type material examined (BMNH). SRI LANKA: 1 ♂ (holotype), “Type” (red circle, printed), “Sarima cretatus Dist type” (handwritten in ink), “Pundalu-oya. Ceylon (printed), 5–03 (handwritten in pencil)”, “1617” (handwritten in ink), “Distant Coll. 1911–383.” (printed). Comparison. The specimen examined appears very similar to S. illibata; it differs, however, in much darker coloration, which looks artefactual. Note. According to the original description, DISTANT (1906) was working with a single specimen from Pundalu-oya (central Sri Lanka, Kandy District).

Sarima elongata Melichar, 1903 (Figs 6, 10, 11) Sarima elongata Melichar, 1903: 80.

Type material examined (ZMHB). SRI LANKA: 1 ♂ (lectotype, here designated), “Type” (red, printed), “6188” (printed), “Ceylon Nietner” (handwritten in ink), “elongata (handwritten in ink) det. Melichar. (printed)”, “Sarima elongata Melichar *” (handwritten in ink), “Mus. Berol.” (printed); 1 ♂ (paralectotype), “Type” (red, printed), “Cat No. 6188” (handwritten in ink), “Ceylon Nietner” (handwritten in ink), “elongata (handwritten in ink) det. Melichar. (printed)”, “Sarima elongata Melichar *” (handwritten in ink), “Mus. Berol.” (printed); 1 ♀ (paralectotype), “Type” (red, printed), “Cat No. 6188” (handwritten in ink), “Ceylon Nietner” (handwritten in ink), “Sarima elongata Melichar *” (handwritten in ink), “elongata (handwritten in ink) det. Melichar. (printed)”, “Mus. Berol.” (printed). Supplementary description. Pro- and mesonotum with no median carina or with very weak carina. Forewings with narrow hypocostal plate. General coloration light yellow- greenish. Forewings with vague light brown patches between median and cubitus anterior in distal half of the wing and on clavus. Males sometimes with 3 whitish spots between median and cubitus anterior distally and with one whitish spot in distal part of clavus. Hind wings matt, with light brown veins. Leg spines black. Total length: males – 5.7 mm; female – 6.0 mm.

178 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Notes on Sarima with description of new genus

Figs 3–4. Forewing in lateral view: 3 – Sarima illibata Melichar, male lectotype; 4 – Pavelauterum fusculum (Melichar), female lectotype. RL –loop of anterior branch of radius.

Note. MELICHAR (1903) mentioned three specimens deposited in the Museum für Naturkunde (Berlin) as the type series. Here I designate the lectotype for a male according to the ICZN (1999: Art. 74) to stabilize the nomenclature in the genus.

Genus Pavelauterum gen. nov. Type species: Hysteropterum fusculum Melichar, 1903. Description. Metope quite narrow, slightly convex, weakly enlarged above clypeus, with distinct median carina running from its upper margin to metopoclypeal suture, and no sublateral carinae (Figs 2, 12). Upper margin of metope weakly concave. Postclypeus

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 179 V. M. GNEZDILOV slightly flattened dorso-ventrally, with no carina. Ocelli present. Pedicel nearly spherical. Coryphe transverse, twice as wide as long at centre, anterior margin weakly convex, posterior margin obtusely angulate (Fig. 13). Pronotum with anterior margin right- angular, keel-shaped and elevated, posterior margin almost straight. Paradiscal fields of pronotum very narrow behind eyes. Paranotal lobes of pronotum wide and rounded, flat, with no carina. Mesonotum slightly longer than pronotum, with weak median carina and distinct lateral carinae. Forewings quite wide, widely rounded (almost truncate) apically (Figs 4, 7), without hypocostal plate. Basal cell oval, precostal area with transverse veins. Radius bifurcate (dividing near basal cell), median with 4 branches (dividing in distal half of wing), cubitus anterior bifurcate (dividing near wing mid-point); many secondary transverse veins present in most cells (Fig. 4). Clavus as long as nearly 4/5 of wing length, open (cubitus posterior and postcubitus + first anal vein joint apically); postcubitus and first anal vein joining at mid-point of clavus. Hind wings possibly three- lobed (feature not readily visible), nearly as long as forewings. Hind tibia with two lateral spines distally and 8 apical spines. First metatarsomere with two latero-apical and 7 intermediate apical spines forming a complete, arch-shaped row. Female sternum VII with hind margin weakly convex. Gonoplacs convex, rounded. Female anal tube long and narrow, rounded apically, anal column short. Comparison. The new genus differs from Sarima in the absence of sublateral carinae from the metope (Figs 2, 12), wider forewings without hypocostal plate and without the unique loop in anterior branch of radius (Figs 4, 7). Externally the genus is similar to Darwallia Gnezdilov, 2010 which also has the metope without sublateral carinae and forewings widely rounded apically with median vein dividing into 4 branches. However, Darwallia is distinguished by its elongate coryphe and carinate clypeus (GNEZDILOV 2010).

Pavelauterum fusculum (Melichar, 1903) comb.nov. (Figs 2, 4, 7, 12, 13) Hysteropterum fusculum Melichar, 1903: 77. Sarima fuscula, MELICHAR (1906): 302.

Type material examined (ZMHB). SRI LANKA: 1 ♀ (lectotype, here designated), “Type” (red, printed), “Ceylon Nietn.” (yellow, handwritten in ink), “fusculum” (handwritten in ink), “Hysteropterum fusculum Melichar *” (handwritten in ink), “Mus. Berol.” (printed); 1 ♀ (paralectotype), “Type” (red, printed), “Cat. No. 7034” (handwritten in ink), “Ceylon Nietner” (handwritten in ink), “fusculum (handwritten in ink) det. Melichar. (printed)”, “Hysteropterum fusculum Melichar *” (handwritten in ink), “Mus. Berol.” (printed), “Sarima fuscula” (handwritten in ink). Supplementary description. General coloration light brown. Hind wings matt, with dark brown veins. Leg spines black. Total length: females – 4.7 mm. Note. MELICHAR (1903) mentioned two specimens deposited in the Museum für Naturkunde (Berlin) as the type series. Here I designate a female lectotype according to the ICZN (1999: Art. 74) to stabilize the nomenclature.

180 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Notes on Sarima with description of new genus

Figs 5–13. Habitus, lateral view: 5 – Sarima illibata Melichar, female paralectotype; 6 – Sarima elongata Melichar, male lectotype; 7 – Pavelauterum fusculum (Melichar), female lectotype. Head, frontal (8, 10, 12) and dorsal (9, 11, 13) view: 8–9 – Sarima illibata Melichar, female paralectotype; 10–11 – Sarima elongata Melichar, male lectotype; 12–13 – Pavelauterum fusculum (Melichar), female lectotype. Scale bar: 1 mm.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 181 V. M. GNEZDILOV

Acknowledgements My sincere thanks to Dr Igor Malenovský (Brno, Czech Republic), Dr Jürgen Deckert (Berlin, Germany), Mr Mick Webb and Dr David Ouvrard (London, United Kingdom), and Prof. Dr Thierry Bourgoin (Paris, France) who kindly made material available for study, Dr Masami Hayashi (Saitama, Japan) and Dr Ursula Göllner- Scheiding (Berlin, Germany) for their help with literature, and Prof. Dr Hannelore Hoch (Berlin, Germany) for her hospitality in the Museum für Naturkunde. The study was financially supported by the Alexander von Humboldt Stiftung (Bonn, Germany).

References

ANUFRIEV G. A. & EMELJANOV A. F. 1988: Podotryad Cicadinea (Auchenorrhyncha). [Suborder Cicadinea (Auchenorrhyncha).] Pp. 12–495. In: LER P. A. (ed.): Opredelitel’ nasekomykh Dal’nego Vostoka SSSR v shesti tomakh. Vol. 2. Ravnokrylye i poluzhestkokrylye. [Keys to the insects of the Far East of the USSR in six volumes. Volume II Homoptera and Heteroptera.] Nauka, Leningrad, 972 pp (in Russian). BOURGOIN T. 2012: FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha. Version 8, updated 30 June 2013. Available online at http://flow.snv.jussieu.fr/ (last accessed on 1 July 2013). CHAN M. L. & YANG C. T. 1994: Issidae of Taiwan (Homoptera: Fulgoroidea). Chen Chung Book, Taichung, 188 pp. DISTANT W. L. 1906: Rhynchota. – Vol. III (Heteroptera–Homoptera). The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, 503 pp. EMELJANOV A. F. 1971: Novye rody tsikadovykh fauny SSSR iz semeystv Cixiidae i Issidae (Homoptera, Auchenorrhyncha). (New genera of planthoppers of the families Cixiidae and Issidae (Homoptera, Auchenorrhyncha) from the fauna of the USSR.) Entomologicheskoe Obozrenie 50: 619–627 (in Russian with English summary; English translation published in Entomological Review 50: 350–354). EMELJANOV A. F. 2001: Larval characters and their ontogenetic development in Fulgoroidea (Homoptera, Cicadina). Zoosystematica Rossica 9: 101–121. GNEZDILOV V. M. 2010: Three new genera and three new species of the family Issidae (Hemiptera: Fulgoromorpha) from Borneo and Sumatra. Tijdschrift voor Entomologie 153: 41–52. HORI Y. 1970: Genus Sarima Melichar of Japan, with the description of a new Ryukyu species (Hemiptera: Issidae). Transactions of Shikoku Entomological Society 10(3–4): 79–83. HORI Y. 1971: Notes on some Philippine Issidae (Hemiptera). Transactions of Shikoku Entomological Society 11(2): 60–70. ICZN (International Commission on Zoological Nomenclature) 1999: International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature c/o the Natural History Museum, London, XXIX + 306 pp. MELICHAR L. 1903: Homopteren-fauna von Ceylon. Verlag von Felix L. Dames, Berlin, 248 pp. MELICHAR L. 1906: Monographie der Issiden (Homoptera). Abhandlungen der K. K. Zoologisch-botanischen Gesellschaft in Wien 3(4): 1–327. METCALF Z. P. 1958: General catalogue of the Homoptera. Fasc. IV. Fulgoroidea. Part 15. Issidae. Waverly Press, Baltimore, 561 pp.

182 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 183–189, 2013

Two new species and additional records of the genus Eupteryx (Hemiptera: Cicadellidae: Typhlocybinae) from the northern Caucasus

ROLAND MÜHLETHALER1 & VLADIMIR M. GNEZDILOV2 1 Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, 10155 Berlin, Germany; e-mail: [email protected] 2 Zoological Institute, Russian Academy of Sciences, Universitetskaya nab.1, 199034 Saint Petersburg, Russia; e-mail: [email protected]

MÜHLETHALER R. & GNEZDILOV V. M. 2013: Two new species and additional records of the genus Eupteryx (Hemiptera: Cicadellidae: Typhlocybinae) from the northern Caucasus. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 183–189. – Two new species of the genus Eupteryx Curtis, 1833 (E. lautereri sp.nov. and E. logvinenkoae sp.nov.) are described from the mountain regions of the Krasnodar Territory in Russia and Abkhazia in Georgia. Eupteryx filicum (Newman, 1853) is recorded for the first time for Russia and Eupteryx calcarata Ossiannilsson, 1936 for Georgia. Keywords. Auchenorrhyncha, Cicadomorpha, , faunistics, taxonomy, new species, Russia, Krasnodar Territory, Georgia, Abkhazia

Introduction The genus Eupteryx Curtis, 1833 includes two subgenera Eupteryx (sensu stricto) and Stacla Dworakowska, 1969 and 116 species to date (ASCHE & HOCH 2004, GUGLIELMINO et al. 2011, POGGI 2012, DMITRIEV & DIETRICH 2003), most of which are distributed in the Palaearctic region (78 species). Several species-groups were proposed by RIBAUT (1936) mainly based on wing venation and adult genital characters, later supported by STEWART (1986, 1988) using larval characters. In Europe and North America, some species of Eupteryx are important pests of medical and culinary herbs in terms of feeding damage (HENKE et al. 2013). Here we describe two new Eupteryx (s. str.) species from the Krasnodar Territory of Russia and Abkhazia in Georgia. We also provide some new faunistic information on two species of the genus from the northern Caucasus. We are happy to take the opportunity to dedicate one of the species described below to RNDr. Pavel Lauterer on the occasion of his 80th birthday and in honour of his enormous contribution to knowledge of leafhoppers and psyllids of Europe.

Material and methods The specimens examined are deposited in the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (ZIN) and the Moravian Museum, Brno, Czech Republic (MMBC).

183 R. MÜHLETHALER & V. M. GNEZDILOV

Photographs of the specimens were taken using a Leica MZ8 microscope with a JVC KY F7OB video camera. The images were processed with Synoptics Automontage software. Drawings of the internal genital characters were made from temporary slide- mounts in glycerine-gelatine using a camera lucida.

Taxonomy Eupteryx lautereri sp.nov. (Figs 1–7, 12–15)

Type material. Holotype: ♂, Russia, Krasnodar Territory: Aibga Range, 1900 m, 8 km SE of Krasnaya Polyana, 11.vii.1999, V. M. Gnezdilov leg. (ZIN). Paratypes: Russia, Krasnodar Territory: 1 ♂, 1 ♀, same data as holotype (ZIN). Georgia, Abkhazia: 2 ♂♂ 5 ♀♀, 1 intersex, Bzybsky Range, 13–18.viii.1931, Voronov leg. (ZIN: 2 ♂♂, 4 ♀♀, 1 intersex; MMBC: 1 ♀); 1 ♂ (parasitized), Gudaut, 3–4.viii.1931, Voronov leg. (ZIN). Description. Body length: males: 3.9–4.1 mm; females: 3.7–4.0 mm; intersex: 3.7 mm. Coloration. Face yellowish-greenish, almost without dark markings (Figs 13, 15). Vertex with two distinct, round, black-brown spots, sometimes weak or absent, not reaching fore-margin of head in dorsal view (Figs 12, 14). Antennal pit often with black triangular spot below eye. Pronotum yellowish-greenish without dark markings. Scutellum with two large, dark lateral triangles, sometimes weak or absent (Figs 12, 14). Forewing with four distinct black-brown spots, two along commissural border, one along radial vein (in its two-thirds), one in median cell. Apical cells of forewing fuscous (Figs 12, 14). Male genitalia. Pygofer process long and curved (Fig. 1). Appendages of aedeagus bent inwards with tips crossing each other dorsal to the shaft (Figs 3–5). Genital style as in Fig. 6. Etymology. The species is dedicated to RNDr. Pavel Lauterer. Differential diagnosis. Eupteryx lautereri sp.nov. belongs to the E. aurata species- group. The shape of the aedeagus is similar to that of many species of the E. aurata species group. The pygofer process is similar to E. fahringeri Melichar, 1911 (see DLABOLA 1981 for characters of the male). Very pale-coloured specimens with almost no markings on the forewings look similar to E. lelievrei Lethierry, 1874. The song apodemes are relatively long, reaching almost to the fourth abdominal segment (Fig. 7). The combination of the coloration and the pygofer process is unique and separates E. lautereri sp.nov. clearly from all other Eupteryx spp. Ecology. In Krasnodar Territory the species was collected in subalpine meadows (1900 m) by sweeping herbs. Note. The species was erroneously recorded from Russia as “Eupteryx ?lelievrei (Lethierry)” by GNEZDILOV (2000).

Eupteryx logvinenkoae sp.nov. (Figs 8–11, 16–17)

Type material. Holotype: ♂, Russia, Krasnodar Territory: near Novorossijsk, 25.vii.1997, V. M. Gnezdilov leg. (ZIN). Paratypes: 2 ♀♀, Russia, Krasnodar Territory: near Gelendzhik, 21.vii.1997, V. M. Gnezdilov leg. (ZIN).

184 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Eupteryx

Figs 1–7. Eupteryx lautereri sp.nov. 1 – male genital segment in lateral view; 2 – male genital segment in ventral view; 3 – aedeagus in lateral view; 4 – aedeagus in dorsal view; 5 – aedeagus in ventral view; 6 – left genital style; 7 – abdominal segments 1–4 and song apodemes in ventral view. Scale bars: 0.2 mm (1, 2, 7); 0.1 mm (3–6).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 185 R. MÜHLETHALER & V. M. GNEZDILOV

Figs 8–11. Eupteryx logvinenkoae sp.nov. 8 – male genital segment in lateral view; 9 – aedeagus in dorsal view; 10 – aedeagus in lateral view; 11 – right genital style. Scale bars: 0.2 mm (8), 0.1 mm (9–11).

Description. Body length: males: 2.5 mm; females: 2.6–2.8 mm. Coloration. Face greenish with distinct black markings (Fig. 17). Two black spots between eyes; two round, black spots on transition to vertex. Genae on each side of frontoclypeus with black band extending to antennal pit. Anteclypeus black (Fig. 17). Vertex in dorsal view with three distinct black spots, two round spots on transition to face, one triangular spot on posterior margin of head. Pronotum with distinct brownish-black markings (Fig. 16). Scutellum with four black markings, two triangular spots at anterior margin, two small, round spots medially. Forewing with greenish-brown markings (Fig. 16). Male genitalia. Pygofer process relatively short and slightly curved (Fig. 8). Appendages of aedeagus evenly bent, not (or only slightly) crossing each other dorsal to shaft (Figs 9–10). Genital style as in Fig. 11.

186 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Eupteryx

Figs 12–17. 12–15 – Eupteryx lautereri sp.nov., habitus: 12 – female, dorsal view; 13 – female, frontal view; 14 – male, dorsal view; 15 – male, frontal view. 16–17 – Eupteryx logvinenkoae sp.nov.: 16 – female, dorsal view; 17 – female, frontal view. Scale bars: 1.0 mm.

Differential diagnosis. Eupteryx logvinenkoae sp.nov. belongs to the E. melissae species-group. The shape of the aedeagus is similar to that of E. melissae Curtis, 1837, E. cytinsularis Guglielmino, Lauterer et Bückle, 2011 and E. ichnusae Poggi, 2012 but the apical appendages do not cross each other and are bent dorsal to the shaft. The structure of the aedeagus is therefore unique and a clear diagnostic character. The pygofer is also similar to that of E. melissae, but the postero-dorsal angle is less prominent (Fig. 8). The coloration of the head and wings is similar to other species of the E. melissae species-group, e.g. E. collina (Flor, 1861). Ecology. The species was collected by sweeping herbs in the sub-Mediterranean plant communities of the Markotkh Range, near the Black Sea coast. Etymology. The species is named in honour of Dr. Valentina Nikolaevna Logvinenko (1929–1983) who studied the Auchenorrhyncha fauna of the Caucasus closely.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 187 R. MÜHLETHALER & V. M. GNEZDILOV

New records Eupteryx calcarata Ossiannilsson, 1936

Material examined. 1 ♂, Georgia, Abkhazia: Kheknara, 500 m, 31.viii.1976, A. K. Zagulyaev leg. (ZIN). Note. A western Palaearctic species, widely distributed in Europe and Kazakhstan (NAST 1972). First record from Georgia.

Eupteryx filicum (Newman, 1853)

Material examined. 2 ♂♂, 3 ♀♀, Russia, Krasnodar Territory: near Lazarevskoe village, 27.ix.1985, A. G. Kireychuk leg. (ZIN). Note. The species was incorrectly identified as Zygina rhamni Ferrari, 1882 and accordingly erroneously recorded for Russia (GNEZDILOV 2000). A western Palaearctic species known from western Europe, the Mediterranean region and Turkmenistan (NAST 1972). Now recorded here for the first time for Russia.

Discussion Eupteryx lautereri sp.nov. clearly belongs to the E. aurata species-group, to judge by its wing venation and the structure of the male genitalia (RIBAUT 1936). To date, only two species of the group may be regarded as closely related: E. fahringeri and E. lelievrei. E. logvinenkoae sp.nov. is part of the E. melissae species-group. Recently, two other species of this group were described: E. cytinsularis and E. ichnusae (GUGLIELMINO et al. 2011, POGGI 2012). Both are from the Mediterranean region and are different in the structure of the aedeagus from E. logvinenkoae sp.nov. described here. The list of Eupteryx species known from the Krasnodar Territory of Russia currently comprises 14 species (GNEZDILOV 2000 and new data in this paper): E. adspersa Herrich- Schäffer, 1838; E. assectator Logvinenko, 1966; E. atropunctata (Goeze, 1778); E. cyclops Matsumura, 1906; E. egregia Logvinenko, 1981; E. filicum; E. florida Ribaut, 1936; E. formaster Logvinenko, 1966; E. lautereri sp.nov.; E. logvinenkoae sp.nov.; E. origani Zachvatkin, 1948; E. praestabilis Logvinenko, 1966; E. stachydearum (Hardy, 1850); and E. tenella (Fallén, 1806). The same pattern of montane distribution (on the Aibga and Bzybsky Ranges – both of them parts of the Main Caucasus Range) appears for E. lautereri sp.nov. and Mycterodus aspernatus Gnezdilov, 2001 (Issidae) (GNEZDILOV 2001). Apparently both species are north-western Caucasian endemics occurring in subalpine meadows. As well as E. lautereri sp.nov., three more Eupteryx species have been recorded from subalpine meadows in the Aibga Range – E. assectator, E. atropunctata, and E. florida (GNEZDILOV 2000). The intersex of E. lautereri sp.nov. (i.e. a feminized male) was possibly the result of a Wolbachia infection; similar intersexes have recently been reported for other Eupteryx populations (HENKE et al. 2013).

188 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Eupteryx

Acknowledgements We are sincerely grateful to Dr Irena Dworakowska (Vancouver, Canada) for her comments at the initial stage of identification of the materials, to Dr Valerio Mazzoni (San Michele all’Adige, Pisa, Italy) for his identification of Eupteryx filicum, to Prof. Dr Georgij A. Anufriev (Nizhnij Novgorod, Russia) for his comments on distribution of Eupteryx species, and Dr Aleksandr A. Stekol’nikov (Saint Petersburg, Russia) and Dr Valerij Shchyurov (Krasnodar, Russia) for their company during field trips with V. M. Gnezdilov to Krasnodar Territory. Finally we would like to thank Dr Dmitry Dmitriev (Champaign, Illinois, USA) for his helpful notes on the manuscript. V. M. Gnezdilov was financially supported by the Royal Society of London (United Kingdom) and the Alexander von Humboldt Stiftung (Bonn, Germany).

References

ASCHE M. & HOCH H. 2004 onward: Fauna Europaea: Cicadomorpha. In: Fauna Europaea version 1.0, Available online from: http://www.faunaeur.org (accessed on 17 July 2013) DLABOLA J. 1981: Ergebnisse der Tschechoslowakisch-Iranischen Entomologischen Expeditionen nach dem Iran (1970 und 1973). (Mit Angaben über einige Sammelresultate in Anatolien). Homoptera: Auchenorrhyncha (II. Teil). Acta Entomologica Musei Nationalis Pragae 40: 127–311. DMITRIEV D. & DIETRICH C. 2003 onward: 3i interactive keys and taxonomic databases, subfamily Typhlocybinae. Available online from: http://imperialis.inhs.illinois.edu/dmitriev (accessed on 17 July 2013). GNEZDILOV V. M. 2000: To the knowledge of the faunistic complexes of the Cicadina (Homoptera) in the main plant formations of the Northwestern Caucasus. Entomologicheskoe obozrenie 79(4): 794–811 (in Russian, English translation published in Entomological Review (2001) 80(8): 927–945). GNEZDILOV V. M. 2001: New and little known leafhoppers and planthoppers from Caucasus (Homoptera, Cicadina). Zoosystematica Rossica 9(2): 359–364. GUGLIELMINO A., LAUTERER P. & BÜCKLE C. 2011: Eupteryx cytinsularis, a new species of the melissae group (Rhynchota Auchenorrhyncha Cicadellidae) from Sicily, Sardinia and Corsica. Bulletin of Insectology 64: 23–26. HENKE C., NICKEL H., SCHEU S. & SCHAEFFER I. 2013: Evidence for Wolbachia in leafhoppers of the genus Eupteryx with intersexual morphotypes. Bulletin of Insectology 66(1): 109–118. NAST J. 1972: Palaearctic Auchenorrhyncha (Homoptera). An annotated check list. Polish Scientific Publishers, Warszawa, 550 pp. POGGI F. 2012: Descrizione di due nuove specie Italiane di Typhlocybinae (Hemiptera, Cicadellidae). Doriana 8(373): 1–8. RIBAUT H. 1936: Homoptères Auchénorhynques. I. (Typhlocybidae). Faune de France 31. Paul Lechevalier et Fils, Paris, 231 pp. STEWART A. J. A. 1986: Descriptions and key to the nymphs of Eupteryx (Curtis) leafhoppers (Homoptera: Auchenorrhyncha) occurring in Britain. Systematic Entomology 11: 365–376. STEWART A. J. A. 1988: Patterns of host-plant utilization by leafhoppers in the genus Eupteryx (Hemiptera: Cicadellidae) in Britain. Journal of Natural History 22: 357–379.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 189

ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 191–228, 2013

New species of Docalidia from Peru with new records (Hemiptera: Cicadomorpha: Cicadellidae: Coelidiinae)

MERVIN W. NIELSON1 & PEDRO W. LOZADA2 1 Retired, former Affiliate Faculty, Monte L. Bean Museum, Brigham Young University, Provo, UT 84602, USA; e-mail: [email protected] 2 Departamento de Entomologia, Museo de Historia Natural, UNMSM, Av. Arenales, Apartado 14-0434, Lima 14, Peru; e-mail: [email protected]

NIELSON M. W. & LOZADA P. W. 2013: New species of Docalidia from Peru with new records (Hemiptera: Cicadomorpha: Cicadellidae: Coelidiinae). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 191–228. – The following 19 new species of Docalidia Nielson, 1979 (Hemiptera: Cicadomorpha: Cicadellidae: Coelidiinae: Teruliini) from Peru are described, illustrated and photographed: Docalidia acucifera sp.nov., D. basispinosa sp.nov., D. bilobata sp.nov., D. collaris sp.nov., D. crassitudinis sp.nov., D. hama sp.nov., D. inuncta sp.nov., D. lineolaris sp.nov., D. maldonadoi sp.nov., D. medispinata sp.nov., D. minatura sp.nov., D. minima sp.nov., D. pakitzaensis sp.nov., D. parabistyla sp.nov., D. parainsolita sp.nov., D. pedalis sp.nov., D. reticulata sp.nov. D. setosa, sp.nov. and D. trulla sp.nov. Docalidia dentatula (Metcalf, 1964) and D. zanolae Nielson, 1986 are new records for Peru. Keywords. Auchenorrhyncha, Cicadellidae, Coelidiinae, Teruliini, Leafhoppers, Docalidia, taxonomy, new species, new records, Neotropical Region, Peru

Introduction The genus Docalidia Nielson, 1979 is the largest group of coelidiine leafhoppers in the New World, known only from the Neotropical region northward to Honduras and southward to Argentina. Only one species is known in each of these countries. Since the genus was first described, 179 species have been elucidated (NIELSON 1979, 1982a, 1982b, 1982c, 1986, 1990, 1992, 1996, 2011). Key to species is provided in NIELSON (1979, 1986, 2011). In this paper 19 new species are added which now comprise 195 known species in the genus. Among these, 78 have been recorded from Peru and all are catalogued in NIELSON (2011), except 2 (new records) and 19 described herein. Peru has the richest fauna followed by Brazil, Colombia and Bolivia. Species occupying more than one country is very low; only 10.7% among 16 countries in its range. Species diversity is very low in the extreme southern and northern ranges of the Neotropical region.

Material and methods Specimens for this study were provided entirely by the junior author who collected much of the material. The material is deposited in the following institutions: Departamento de Entomologia, Museo de Historia, Lima, Peru (UNMSM) and Monte L. Bean Museum, Brigham Young, University, Provo, UT 84602, Utah, USA (MLBM).

191 M. W. NIELSON & P. W. LOZADA

Taxonomy Genus Docalidia Nielson, 1979 Docalidia Nielson 1979: 179. Type species: Jassus ruficosta Jacobi 1905, by original designation. Diagnosis. Members of the genus are described by the following combination of features: small to large size, robust form; color concolorous to well marked; head short, broad. often about as wide as pronotum; pronotum and mesonotum large; forewings often broad throughout; male pygofer large, often with short to long, caudoventral process, sometimes with caudodorsal margin produced to long process; segment X often with ventral process, exposed or cryptic; aedeagus always long, very slender, tubular, with single, short to long, subapical process on ventral margin, process sometimes absent; style slender to robust, apical half to one third often large, often with numerous, secondary processes in apical half; connective T-shaped, stem short to long, configuration variable; dorsal connective short, very narrow; subgenital plate large, broad, rarely narrow, glabrous to setaceous. Basic features that separate the genus from other teruliine genera of the New World are the following combination of characters: robust form, the long, very narrow shaft of the aedeagus with a single subapical process arising from the ventral margin of the shaft (sometimes broken off or absent) and the very broad, subgenital plate.

Description of species Docalidia acucifera sp.nov. (Figs 1–7, 134)

Type material. Holotype ♂: PERU, Castilla, 345 m., 10°10′S 75°15′W, 11.vi.87, P. Lozada (UNMSM). Paratype, 1 ♀, same data as holotype (UNMSM). Description. Length. Male 6.80 mm, female 7.70 mm. External morphology. Medium-sized, robust species. General color dark brown to black; head light brown with black markings on crown in male, crown light tan in female; face black in male, brown in female. Head large, narrower than pronotum, anterior margin broadly rounded; crown broad, about as wide as eyes, short, slightly wider than long, slightly produced anteriorly, lateral margins convergent basally; eyes large, semiglobular; clypeus long, lateral margins sinuate posteriorly, clypellus long, narrow, slightly tumid basally. Male genitalia. Pygofer in lateral view large, with long, fairly robust caudodorsal process, caudoventral process long, very narrow, sharply pointed (Fig. 1); segment X with prominent ventral process, very broad in basal half, curved dorsally, narrowed and sharply pointed in apical half (Fig. 1); aedeagus, shaft very long, very narrow in dorsal and lateral views, without subapical process (Figs 2, 3); style very long, robust in dorsal view, narrow in lateral view except inflated in distal 1/3 with narrow band of dense microsetae on lateral margin, small curved spine medially on lateral margin (Figs 4, 5); dorsal connective short, narrow (Fig. 3); connective large, T-shaped (Fig. 6); subgenital plate long, broad, glabrous (Fig. 7).

192 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Etymology. The name of this species is descriptive for acutely narrow aedeagal shaft. Remarks. From D. basispinosa sp.nov. to which it is similar in aedeagal and stylar configurations, D. acucifera sp.nov. is easily distinguished by the narrow ventral process of segment X, presence of medial spine on the style and broad basal half of the caudoventral pygofer process.

Docalidia basispinosa sp.nov. (Figs 8–14, 135)

Type material. Holotype ♂: PERU: LO [Loreto], Requena, Jenaro Herrera, 73°40′W, 4°55′S, 21.viii.1990, P. Lozada (UNMSM). Description. Length. Male 6.10 mm, female unknown. External morphology. Small, robust species. General color black throughout except head; crown light yellow with small markings; eyes light green tinged with light brown; face black except for narrow light yellow on anterior margin. Head small, narrower than pronotum, anterior margin broadly rounded; crown short, broad, about as wide as width of eyes, produced beyond anterior margin of eyes, anterior margin obtusely angled, disk depressed on each side of middle, lateral margins slightly convergent basally; eyes large, elongate ovoid; clypeus long, broad, lateral margins broadly convex; clypellus broad. Male genitalia. Pygofer in lateral view narrow, elongate, glabrous, caudodorsal process very long, nearly as long as dorsal margin of pygofer, sharply triangulate apically, caudoventral process very broad basally, narrow sharp spine apically (Fig. 8); segment X with long, narrow ventral process, sharply pointed in 1/3 (Fig. 8); aedeagus long, shaft very narrow, broadly curved in lateral view, very short process subapically (Figs 9, 10); style long, apophysis rectangulate in near distal half in dorsal view, row of short, dense setae compacted on lateral margin, few setae apically (Figs 11, 12); dorsal connective short, plate-like (Fig. 9); connective large, arms narrow, membrane and medial ridge absent, stem large, subquadrate (Fig. 13); subgenital plate long, very broad (Fig. 14). Etymology. The name of this species is descriptive for the short row of small spines on the ventral base on the pygofer caudodorsal process. Remarks. This species belongs to a large group in which the style is rectangulate in near distal half with a compact lateral row of short setae. From these species, D. basispinosa sp.nov. can be separated easily by the row of short ventral spines at base of caudodorsal pygofer process.

Docalidia bilobata sp.nov. (Figs 15–21, 136)

Type material. Holotype ♂: PERU: MD [Madre de Dios], 15 km. E. Puerto Maldonado, 200 m., 4.ii.1990, P. Lozada (UNMSM). Paratype ♂: PERU: MD [Madre de Dios], Z.R. [Zona Reservada], Tambopata, 290 m, 19.iii.87, P. Lozada (MLBM). Description. Length. Male 7.70 mm, female unknown. External morphology. Medium-sized, robust species. General color light brown with numerous black markings; forewings light brown with numerous black markings in basal

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 193 M. W. NIELSON & P. W. LOZADA half, veins interspersed with small black and light brown spots; mesonotum and pronotum light brown, speckled with black markings; crown light brown; eyes light brown suffused with dark brown along posterior margin; face light brown with numerous black markings; clypeus light brown, row of short, transverse black stripes on each side, coalesced medially, clypellus light brown, genae and lorae black bordered by light brown. Head large, short, narrower than width of eyes, anterior margin broadly rounded; crown short, about as long as wide, narrower than width of eyes; eyes large, semiglobular; clypeus long broad anteriorly, lateral margins broadly convex; clypellus narrow. Male genitalia. Pygofer in lateral view with very long caudodorsal process, small digitate caudoventral lobe, glabrous (Fig. 15); segment X ventral process absent; aedeagus long, very narrow, tubular, with long subapical process on ventral margin, small flange medially (Figs 16, 17); style very long, robust, inflated in distal 1/5, in dorsal view with two visible lobes in distal 1/5, one lobe glabrous, bisected, one profusely and densely setaceous (Figs 18, 19); dorsal connective short, strap-shaped (Figs 16, 17); connective large, lateral arms broad, long, membrane and medial ridge absent, stem bifurcate (Fig. 20); subgenital plate long, very broad, glabrous (Fig. 21). Etymology. The name of the species is descriptive for the apical, bilobed style. Remarks. This species is unique by virtue of two apical lobes on the style in combination with the bifurcate stem on the connective which will separate the species from all others.

Docalidia collaris sp.nov. (Figs 22–28, 137)

Type material. Holotype ♂: PERU: LO [Loreto], Centro de Investigacion, Jenaro Herrera, 73°45′W 4°55′S, 25.x.1989, P. Lozada (UNMSM). Description. Length. Male 7.90 mm, female unknown. External morphology. Moderately large, very robust species. General color black throughout, face black. Head small, broad, narrower than pronotum, anterior margin broadly rounded; crown very broad, wider than width of eyes, slightly produced anteriorly, lateral margins nearly parallel; eyes large, elongate ovoid; clypeus long, broad, lateral margins broadly convex, clypellus broad, not as wide as clypeus. Male genitalia. Pygofer in lateral view large, caudodorsal process long, pointed apically, caudodorsal process rectangulate, obliquely truncate apically (Fig. 22); segment X with very long ventral process, apical half curved, sharply pointed (Fig. 22); aedeagus short, shaft uniformly wide except apically, in ventral view with median, longitudinal ridge to apex, ventral subapical process absent (Figs 23, 24); style long, longer than aedeagus, moderately robust, with long process below middle on lateral margin in dorsal view (Figs 25, 26); dorsal connective moderately long, strap-shaped (Fig. 23, 24); connective large, arms horizontal, membrane and medial ridge absent, stem semiovate (Figs 27); subgenital plate large, very broad, profusely setose (Fig. 28). Etymology. The name of the species is descriptive for the ventral longitudinal ridge (carina) on the aedeagal shaft.

194 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Remarks. The following combination of distinctive characters will separate D. collaris sp.nov. from all species: aedeagus glabrous with ventral, longitudinal median ridge, style with digitate process distad of middle, subgenital plate with numerous long microsetae, segment X with long, narrow ventral process.

Docalidia crassitudinis sp.nov. (Figs 29–35, 138)

Type material. Holotype ♂: PERU: LO [Loreto], Requena, 4.5 Km. Carretera Jenaro, Herrera- Puerto Angamos, 31.viii.1990, P. Lozada (UNMSM). Additional material examined. 1 ♂ (damaged), same data as holotype (MLBM). Description. Length. Male 8.50 mm, female unknown. External morphology. Moderately large, robust species. General color dark brown to black with small, yellow spots on veins of forewings and pronotum; mesonotum black; crown light brown with black markings; eyes light brown; face black. Head small, narrower than pronotum, anterior margin broadly rounded; crown broad, about as wide as width of eyes, slightly produced anteriorly, lateral margins convex; eyes large, semiglobular; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus. Male genitalia. Pygofer in lateral view narrow, glabrous with moderately long caudodorsal process, small, digitate caudoventral process (Fig. 29); segment X with small, digitate ventral process (Fig. 29); aedeagus long, broad in dorsal view, slightly constricted near middle, narrow, tubular in lateral view with long, subapical process on ventral margin (Figs 30, 31); style long, broad, in dorsal view with row of coarse spines submedially on lateral margin, narrow parallel stripe of dense setae surpassing spines, tuft of fine microsetae in apical 1/6 (Figs 32, 33); dorsal connective short, strap-shaped (Fig. 31); connective large, T-shaped, arms horizontal, stem large, digitate, short, basolateral process on each side (Fig. 34); subgenital plate long, very broad, glabrous (Fig. 35). Etymology. The name of the species is descriptive compound name for the dense, dentate processes on the style. Remarks. From D. pedalis sp.nov. to which it has similar features of the pygofer and style, D. crassitudinis sp.nov. can be distinguished by the broader subgenital plate, presence of the subapical aedeagal process and by dentate processes in combination with microsetae on the style

Docalidia hama sp.nov. (Figs 36–42, 139)

Type material. Holotype ♂: PERU: LO [Loreto], Requena Jenaro Herrera, 73°40′W 4°55′S, 21.viii.1990, P. Lozada (UNMSM). Description. Length. 7.60 mm, female unknown. External morphology. General color dark brown to black with light brown markings; forewings dark brown, narrow uneven broken transverse light brown band below apex of

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 195 M. W. NIELSON & P. W. LOZADA mesonotum, large triangular, light brown marking near middle of costa, with many small spots and irregular shape, light brown markings; mesonotum black; pronotum black with few light brown spots; crown light tan with black markings; eyes tan; face black; clypeus with anterior margin tan. Head large, broad, narrower than pronotum, anterior margin broadly rounded; crown short, broad, wider than width of eyes, slightly produced anteriorly, lateral margins slightly convergent basally; clypeus long, broad, lateral margins broadly convex; clypellus about 1/3 as long as clypeus, broad, narrower than clypeus. Male genitalia. Pygofer in lateral view large, triangulate, glabrous, caudodorsal process very long, narrow, slightly constricted along middle, apex obliquely truncate, caudoventral process very short, curved ventrally (Fig. 36); segment X with small ventral process medially (Fig. 36); aedeagus long, very narrow in dorsal view, base twisted, small, near middle lateral flange, in lateral view, shaft broad in basal half, narrow in distal half, apex curved laterally, very small subapical spine on ventral margin (Figs 37, 38); style very long, robust, in lateral view apophysis with prominent lateral process distad of middle, small subapical process, in dorsal view apophysis sinuate, slightly tumid along middle below lateral process, inner margin slightly undulate before narrow apex (Figs 39, 40); dorsal connective short (Fig. 38); connective large, T-shaped, stem long (Fig. 41); subgenital plate long, very broad medially, glabrous (Fig. 42). Etymology. The name of the species (a noun in apposition) is descriptive for the hook- like process on the apophysis of the style. Remarks. From D. parabistyla sp.nov. to which it is similar in configuration of the style, D. hama sp.nov. can be separated by the very short, curved pygofer caudoventral process (very long in D. parabistyla) and by the glabrous subgenital plate (profusely setaceous in D. parabistyla).

Docalidia iuncta sp.nov. (Figs 43–49, 140)

Type material. Holotype ♂: PERU: MD [Madre de Dios], 10 km. Puerto Maldonado, Rio de Dios, 7.IUv.1986, I. Bohorque (UNMSM). Description. Length. Male 6.10 mm, female unknown. External morphology. General color black grading to light brown with numerous yellow spots or markings; forewings dark brown in basal 1/3 grading to light brown in distal 2/3, with numerous small spots on veins; mesonotum black; pronotum black with numerous yellow spots; crown black, ocelli ringed with yellow; eyes dark brown; face black with few yellow spot on clypeus. Head large, broad, anterior margin broadly rounded, narrower than pronotum; crown longer than wide, narrower than width of eyes, slightly produced anteriorly, lateral margins convergent basally; eyes large, semiglobular; clypeus long, broad, lateral margin broadly convex; clypellus broad, narrower than clypeus. Male genitalia. Pygofer in lateral view small, glabrous, caudodorsal process deeply bifurcate, caudoventral process long, lobe-like (Fig 43); segment X with very broad

196 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia ventral process, small curved spine apically (Fig. 43); aedeagus long, tubular, slender with very small, ventral subapical process (Figs 44, 45); style very long, longer than aedeagus, in dorsal view with robust, elliptical lobe in distal 2/5, lobe dentate on inner lateral margin, cleft longitudinally, in lateral view narrow with triangulate lobe distad of middle (Figs 46, 47); dorsal connective long, narrow (Fig. 45); connective T-shaped, stem small (Fig. 48); subgenital plate short, broad, glabrous (Fig. 49). Etymology. The name of the species is descriptive for the deep basal attachment of the pygofer caudodorsal, bifurcate processes. Remarks. Six species are known with a bifurcate, caudodorsal process on the pygofer. From the other five of these species (D. youngi Nielson, 1979; D. pectinata Nielson, 1979; D. bifurcata Nielson, 1979; D. hansoni Nielson, 1982; and D. paradiscissa Nielson, 1990), D. iuncta sp.nov. can be distinguished by the long, elliptical, dentate, apical lobe of the style (rectangulate or without such lobe in other species) and short, digitate, lobe-like caudoventral process (short to long, very narrow caudoventral process in other species).

Docalidia lineolaris sp.nov. (Figs 50–56, 141)

Type material. Holotype ♂: PERU: MD [Madre de Dios], 15 km. E. Puerto Maldonado, 200 m., 30.i.1990, P. Lozada (UNMSM). Description. Length. Male 8.40 mm, female unknown. External morphology. Moderately large, robust species. General color tan with dark brown markings; forewings with large dark brown marking in basal 1/3, large, irregular light brown band medially, large, suffused, light brown marking apically (part of right forewing missing); mesonotum, pronotum, head tan; face light brown. Head moderately sized, narrower than pronotum, anterior margin broadly rounded; crown broad, about as wide as width of eyes, slightly produced medially, disk depressed on each side of middle, lateral margins slightly convergent basally; eyes large, elongate ovoid; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus. Male genitalia. Pygofer in lateral view large, elongate, glabrous, caudodorsal process long, very narrow, caudoventral process extremely long, slightly cleft subapically (Fig. 50); segment X with long, very narrow ventral process (Fig. 50); aedeagus in dorsal and lateral views long, narrow, tubular with narrow flange medially, base of shaft inflated in dorsal view (Figs 51, 52); style extremely long, narrow, with row of short processes in distal 2/7 (Figs 53, 54); dorsal connective short, narrow (Figs 51, 52); connective T- shaped, arms long, stem digitate (Fig. 55); subgenital plate long, very broad, row of short microsetae on inner lateral margin (Fig. 56). Etymology. The name of the species is descriptive for the long, linear aedeagus and style. Remarks. From all other known species, D. lineolaris sp.nov. can be separated easily by the combination of very long, linear features of the aedeagus, style and caudoventral, pygofer process. The row of short setae on style, while not unique, supplements the other characters.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 197 M. W. NIELSON & P. W. LOZADA

Docalidia maldonadoi sp.nov. (Figs 57–63, 142)

Type material. Holotype ♂: PERU: MD [Madre de Dios], 15 km. Puerto Maldonado, 4.ii.1990, 200 m., P. Lozado (UNMSM). Paratype: 1 ♂, same data as holotype except 6.ii.1990 (MLBM). Description. Length. Male 7.40–7.60 mm, female unknown. External morphology. Moderately large, robust species. General color black throughout except for brown head; face black bordered with light tan line; clypeus with spots and short transverse dashes on each side; clypellus, lorae and genae bordered with tan line. Head moderately large, narrower than pronotum, anterior margin broadly rounded; crown broad, about as wide as width of eyes, slightly longer than wide, slightly produced anteriorly, disk slightly depressed, anterior margins nearly parallel; clypeus long, broad, lateral margins broadly convex; clypellus slightly broad, narrower than clypeus, apex flared. Male genitalia. Pygofer in lateral view large, broad, glabrous, caudodorsal process long, stout, segmented in distal 1/3, caudoventral process moderately long, lobe-like (Fig. 57); segment X with short, lobe-like ventral process (Fig. 57); aedeagus short, broad in dorsal view with rounded flange medially on lateral margin, in lateral view very narrow, tubular, without subapical, ventral process (Figs 58, 59); style very long, nearly 1/3 longer than aedeagus, apical third with narrow, compressed band of short setae on outer margins (Figs 60, 61); dorsal connective short, strap-shaped (Figs 58, 59); connective broadly curved with very narrow, lateral arms, stem separated into two short, digitate processes (Fig. 62); subgenital plate long, very broad on outer lateral margin, glabrous (Fig. 63). Etymology. The species is named in honor of Faustino Maldonado, 19th century explorer of the Madre de Dios mountains in Peru. Remarks. The following combination of features will separate D. maldonadoi sp.nov. from all other species: style in lateral view is 1/3 longer than the aedeagus, very slender, bordered by a slender band of compact, short setae in apical 1/6, caudodorsal pygofer process with apical secondary process, aedeagus with medial flange on shaft and the subgenital plate which is very broad medially on the outer lateral margin.

Docalidia mediospinata sp.nov. (Figs 64–70, 143)

Type material. Holotype ♂: PERU: AM [Amazonas], Rio Huahuaga, Camino Imacita-Aramango, 5Vii.1986, I. Bohorquez (UNMSM). Description. Length. Male 8.50 mm, female unknown. External morphology. Moderately large, robust species. General color light to dark brown to black; forewings with clavus and costa light brown, dark brown between; face tan. Head large, narrower than pronotum, anterior margin broadly rounded; crown very broad, wider than width of eyes, produced anteriorly about 1/4 of entire length, lateral margins nearly parallel; eyes large, semiglobular; crown long, broad, lateral margins broadly convex; clypellus narrow, narrower than clypeus, apex flared.

198 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Male genitalia. Pygofer in lateral view small, triangulate, glabrous, caudodorsal process long, narrow, caudoventral process very small, with one seta apically, small lobe below (Fig. 64), segment X with long narrow ventral process, tapered apically (Fig. 64); aedeagus in dorsal view short, very narrow, tubular, sharply tapered to apex, in lateral view curved laterally in apical 1/6, small subapical process on ventral margin (Figs 65, 66); style slightly longer than aedeagus, robust with prominent lateral spine medially, apical 1/4 with row of teeth on lateral margin (Figs 67, 68); dorsal connective short, strap- shaped (Figs 65, 66); connective T-shaped, lateral arms horizontal, membrane and medial ridge absent, stem short, bifurcate apically (Fig. 69); subgenital plate long, broad medially, glabrous (Fig. 70). Etymology. The species name is descriptive for the prominent spine on the middle of stylar apophysis. Remarks. Many species of Docalidia have a prominent medial spine on the style. From them, D. mediospinata sp.nov. is nearest to D. scopa Nielson, 1979 and can be separated by the glabrous subgenital plate, small, setate pygofer caudoventral lobe and the very narrow aedeagal shaft.

Docalidia minatura sp.nov. (Figs 71–77, 144)

Type material. Holotype ♂: PERU: MD [Madre de Dios], P. V. Pakitza, Zona Reservada Manu, 22.ix.1988, 450 m., Malaise trap, H. Blancas (UNMSM). Description. Length. Male 6.80 mm, female unknown. External morphology. Small, very robust species. General color black with numerous, small, irregularly shaped, yellow markings; forewings black, veins well marked with yellow spots, cells with irregularly shaped yellow markings; mesonotum black with yellow mark on each side of middle and near anterior margin; pronotum black, bullae dark yellow; crown mostly yellow with black marking on each side of middle; eyes dark brown; face dark yellow with black and ivory markings; clypeus with short, black transverse stripes on each lateral margin, coalesced medially, eight large ivory spots scattered medially and basally; clypellus light brown, apical margin black; lora yellow with curved black stripe; genae black with inner border yellow. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown broad, wider than long, wider than width of eyes, depressed on each side of middle, lateral margins nearly parallel; eyes large, elongate ovoid; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus, lateral margins parallel. Male genitalia. Pygofer in lateral view small, triangulate, glabrous, caudodorsal process moderately long, narrow, tapered apically, caudoventral process moderately long, asymmetrically bifurcate apically (Fig. 71); segment X with long, narrow ventral process (Fig. 71); aedeagus short, narrow, tubular, in dorsal view tapered to sharp apex, gonopore medial on ventral margin, subapical, ventral process absent (Figs 72, 73); style long, longer than aedeagus, apophysis very slender, without secondary processes (Figs 74, 75); dorsal connective long, narrow (Figs 72, 73); connective small, anterior arms broad,

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 199 M. W. NIELSON & P. W. LOZADA membrane and middle ridge present, stem small, digitate (Fig. 76); subgenital plate short, very broad in apical 4/5, glabrous (Fig. 77). Etymology. The name of the species (a noun in apposition) is descriptive for the small size of the features described and illustrated. Remarks. The combination of the following features will separate D. minatura sp.nov. from all other species: small features, asymmetrical, bifurcate, caudoventral pygofer process, lack of subapical, ventral aedeagal process (atypical in the genus) and glabrous, slender stylar apophysis.

Docalidia minima sp.nov. (Figs 78–84, 145)

Type material. Holotype ♂: PERU: MD [Madre de Dios], Z.R. [Zona Reservada], Tambopata, 290 mm., 14.iii.87, P. Lozada (UNMSM). Description. Length. Male 8.50 mm, female unknown. External morphology. Moderately large, species. General color reddish brown with mesonotum and pronotum mostly black; forewings uniformly reddish brown; mesonotum black with few, deep yellow spots; pronotum black with bullae deep yellow, anterior margin deep yellow; head deep yellow, tinged with dark brown spots on eyes and crown; face black with anterior margin deep yellow, clypellus with deep yellow stripe medially. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown broad, slightly wider than long, wider than width of eyes, slightly produced anteriorly, lateral margins nearly parallel; eyes large, semiglobular; clypeus long, broad, slightly broader anteriorly, lateral margins broadly convex; clypellus broad, narrower than clypeus, apex flared. Male genitalia. Pygofer in lateral view triangulate, glabrous, caudodorsal margin moderately long, with apical digitate process, caudoventral process short, arising as base of caudodorsal process (Fig. 78); segment X with ventral process, truncate apically (Fig. 78); aedeagus in lateral view long, narrow, broadly sinuate in lateral view, with small dentate flange below base of medial process, in ventral view slightly inflated along middle with large opening subbasally in ventral view, moderately long, near medial process (Figs 79, 80); style robust, slightly longer than aedeagus, large triangulate lobe in distal 1/3 small, lateral lobe subapically, in dorsal view, inner lateral margin of lobe with dense, narrow band of microsetae, tuft of setae apically (Figs 81, 82); dorsal connective short, narrow (Fig. 79); connective T-shaped, lateral arms narrow, stem large, elongate (Fig. 83); subgenital plate long, very broad, outer lateral margin broadly convex, few short microsetae apically (Fig. 84). Etymology. The name of the species is descriptive for the small aedeagal flange adorned with few lateral teeth. Remarks. From D. pedalis sp.nov. to which it is similar in style, aedeagus and subgenital plate, D. minima sp.nov. can be distinguished by the much narrower, tubular, sinuate aedeagus with submedial ventral process (absent in D. pedalis), presence of subapical lobe on the style and more robust stem on the connective.

200 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Docalidia pakitzaensis sp.nov. (Figs 85–91, 146)

Type material. Holotype ♂: PERU: MD [Madre de Dios, P.V. Pakitza, Zona Reservada Manu, 24.vi.1988, P. Lozada (UNMSM). Description. Length. Male 8.85 mm, female unknown. External morphology. Moderately large, robust species. General color dark brown to black; forewings dark brown, large pale, brown spot along middle of costa; mesonotum black; pronotum dark brown, narrow dark yellow band anteriorly; crown light brown; eyes dark brown; face black with light brown markings; clypeus black with light row of short transverse stripes on lateral margin, few yellow spots medially, clypellus dark yellow with black stripe on each side of middle; lorae and genae black. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown broad, about as wide as width of eyes, slightly produced anteriorly, depressed medially, lateral margins nearly parallel; eyes large, elongate ovoid; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus, lateral margins parallel to flared apex. Male genitalia. Pygofer in lateral view large, glabrous, caudodorsal process moderately long, segmented subapically, caudoventral process small, digitate (Fig. 85); segment X with moderately long, triangulate ventral process (Fig. 85); aedeagus long, in dorsal view shaft constricted medially and in distal 1/3, sinuate in lateral view, with long, near middle ventral process (Figs 86, 87); style long, longer than aedeagus, broad, in dorsal view apophysis slightly inflated in approximate distal 1/5 with row of short, compacted setae in distal 1/5 on inner lateral margin to apex, continuing basad to near half of length of inflated lobe, small lobe apically (Figs 88, 89); dorsal connective short, strap-shaped (Fig. 86); connective large, T-shaped, lateral arms broad, stem broad basally with two small anterio-lateral lobes, narrowed distally (Fig. 90); subgenital plate long, very large, glabrous (Fig. 91). Etymology. The name of the species is after the Pakitza locality in which several other new coelidiine species have been found. Remarks. This species is very similar to D. maldonadoi sp.nov. in the pygofer and style and can be separated by the much more robust aedeagus with the medial ventral spine (absent in D. maldonadoi), by the very broad arms of the connective and by the larger subgenital plate.

Docalidia parabistyla sp.nov. (Figs 92–98, 147)

Type material. Holotype ♂: PERU: MD [Madre de Dios]; Z.R. [Zona Reservada], Tambopata, 290 m., 21.ii.87, P. Lozada (UNMSM). Description. Length. Male 7.50 mm, female unknown. External morphology. General color dark brown to black with numerous, small yellow markings; forewings dark brown to black, veins with small, yellow spots, cells with small, irregularly shaped, dark yellow markings; mesonotum black with few small yellow spots; pronotum black with numerous small yellow spots; head tannish; face black

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 201 M. W. NIELSON & P. W. LOZADA with yellow markings; clypeus black with row of short, transverse, yellow stripes on each side of lateral margins; clypellus tannish with 2 short, longitudinal black stripes; genae and lorae tannish. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown short, broad, slightly narrower than width of eyes, produced slightly anteriorly, lateral margins nearly parallel, slightly depressed on each side of middle; eyes large, elongate ovoid; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus, lateral margins nearly parallel. Male genitalia. Pygofer in lateral view small, glabrous, caudodorsal process long, narrow; caudoventral process very long, triangulate apically (Fig. 92); segment X with long, narrow, ventral process (Fig. 92); aedeagus in dorsal and lateral views moderately long, inflated basally, shaft very narrow, tubular, with long medial process on ventral margin (Figs 93, 94); style very long, longer than aedeagus, apophysis narrow in lateral view, broad in dorsal view, slightly constricted along middle with very long process near middle (Figs 95, 96); dorsal connective moderately long, narrow (Fig. 93); connective T- shaped, arms horizontal, narrow, with membrane or medial ridge, stem long, digitate (Fig. 97); subgenital plate long, very broad medially, long, microsetae along inner lateral margin (Fig. 98). Etymology. The species name is descriptive for the very long process on the style giving an appearance of two styles and the near relationship to Docalidia bistyla Nielson, 1979. Remarks. From D. bistyla to which it is most closely related, D. parabistyla sp.nov. can be separated by the much narrower aedeagal shaft, presence of segment X ventral process and by the presence of ventral aedeagal process (the two lateral features absent in D. bistyla).

Docalidia parainsolita sp.nov. (Figs 99–105, 148)

Type material. Holotype ♂: PERU: PA [Pasco], Oxapampa, Villa America, 7.x.1987, P. Hocking (UNMSM). Description. Length. Male 6.50 mm, female unknown. External morphology. Small, robust species. General color light brown to black; forewings light to dark brown, veins with light brown spots, narrow, broken, transverse, light brown band medially, large light brown marking near middle of costa; mesonotum black with broad, transverse light brown band medially; pronotum black, bullae light brown; crown yellow; eyes dark brown; clypeus black, light brown anteriorly; clypellus light brown in anterior half, black stripe on each side of middle, black apically. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown broad, slightly produced anteriorly, slightly longer than wide, slightly narrower than width of eyes, lateral margins slightly convergent basally, depressed on each side of middle; eyes large, semiglobular; clypeus long, broad, lateral margins evenly and broadly convex; clypellus broad, much narrower than clypeus, lateral margins parallel. Male genitalia. Pygofer in lateral view large, caudoventral process very long, narrow, broadly curved ventrally, evaginate subapically on ventral margin, caudoventral

202 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia process very short, lobe-like, two short setae between base of processes (Fig. 99); segment X long, very broad (Fig. 99); aedeagus short, narrow tubular with short, subapical process on ventral margin (Figs 100, 101); style very long, nearly twice as long as aedeagus, slightly robust, elongate/triangular in distal 1/5, distinct process subapically (Figs 102, 103); dorsal connective moderately long, narrow (Figs 100, 101); connective with narrow, horizontal arms, without membrane or medial ridge, stem small, apex inflated (Fig. 104); subgenital plate long, very broad in distal 3/5, glabrous (Fig. 105). Etymology. The species name is descriptive for the unusual genitalia features and close relationship to Docalidia insolita Nielson, 1979. Remarks. From D. insolita to which it is similar in pygofer and subgenital plate features, D. parainsolita sp.nov. can be distinguished by the configuration of the apical 1/5 of the style which is elongate/triangulate with one short, subapical lateral spine (one long, curved spine and row of teeth in D. insolita).

Docalidia pedalis sp.nov. (Figs 106–112, 149)

Type material. Holotype ♂: PERU: PA [Pasco], Qbde., Castilla, 345 m., 10°10′S 75°15′W, 11.vi.87, P./ Lozada (Docalidia sp. pusilla ?), P. W. Lozada, 1987 (UNMSM). Description. Length. Male 8.00 mm, female unknown. External morphology. Moderately large, robust species. General color light brown; forewings light brown, mesonotum, pronotum, crown light brown with few, small, dark brown markings; eyes dark brown; face dark brown to black; clypeus with base of antennal sockets black; clypellus, genae and lorae brown. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown broad, nearly as wide as width of eyes, slightly longer than wide, lateral margins nearly parallel, depressed on each side of middle, slightly produced anteriorly; eyes large, semiglobular; clypeus long, broad, lateral margins broadly convex, clypellus broad, narrower that clypeus. Male genitalia. Pygofer in lateral view narrowly triangulate, glabrous; caudodorsal process short; caudoventral process small, lobe-like (Fig. 106); segment X with triangulate, ventral lobe (Fig. 106); aedeagus long, in dorsal view shaft broad in basal 1/4, tapered in distal 3/4, in lateral view with long medial flange on ventral margin, foot- shaped in distal 1/4, ventral subapical process absent (Figs 107, 108); style long, robust, in dorsal view apophysis with triangulate lobe in apical 1/3, inner lateral margin of lobe with long row of short microsetae, in lateral view row of setae very dense in basal half (Figs 109, 110); dorsal connective short, narrow (Figs 107, 108); connective T-shaped, lateral arms broad, horizontal, without membrane and medial ridge, stem short, digitate (Fig. 111); subgenital plate short, broad, with numerous long fine microsetae (Fig. 112). Etymology. The name of the species is descriptive for the aedeagus which is foot-shaped in apical one fourth. Remarks. Several species have a foot-shaped aedeagus. From them, D. pedalis sp.nov. can be separated by the large, setose subgenital plate and elongate pygofer.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 203 M. W. NIELSON & P. W. LOZADA

Docalidia reticulata sp.nov. (Figs 113–119, 150)

Type material. Holotype ♂: PERU: MD [Madre de Dios], Pakitza, Zona Reservada Manu, 22.ix.1988, 450 m., Malaise trap, H. Blancas (UNMSM). Description. Length. Male 6.80 mm, female unknown. External morphology. Small, robust species. General color light yellow to dark brown; forewings dark brown with broad, transverse light yellow bands, one anteriorly below apex of mesonotum, one crossing apex of clavus, veins marked with very small, light yellow spots; mesonotum dark brown with few small yellow markings; pronotum black, mostly covered with light yellow bullae; crown light brown, anterior and posterior margins with dark brown markings; eyes dark brown; clypeus dark brown to black with row of short transverse stripes on each side of lateral margins, coalesced medially, few small, scattered yellow spots; clypellus light brown; lorae and genae yellow with dark brown markings. Head short, broad, narrower than pronotum, anterior margin broadly rounded; crown short, broad, about as wide as width of eyes, slightly produced anteriorly, lateral margins slightly convergent basally, depressed on each side of middle; eyes large, semiglobular; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus, margins parallel. Male genitalia. Pygofer in lateral view small, subquadrate, glabrous, caudodorsal process short, slightly broad, dark, longitudinal reticulations medially; caudoventral process absent (Fig. 113); segment X with moderately long, narrow ventral process (Fig. 113); aedeagus short, in lateral view shaft very broad in basal 2/3, narrowed abruptly and curved in apical 1/3, moderately long, near medial ventral process, in dorsal view shaft narrow, tubular, flange near middle on ventral margin, apex recurved, gonopore medial, exiting laterally (Figs 114, 115); style about as long as aedeagus, apophysis narrow in basal 3/4, very broad, cleaver-shaped lobe in apical 1/4, reticulated markings on lobe (Figs 116, 117); dorsal connective short, narrow in lateral view, base broad in dorsal view (Figs 114, 115), in dorsal view, broad basally (Fig. 115); connective with broad, horizontal arms, membrane and medial ridge absent, stem small, pendulate (Fig. 118); subgenital plate long, moderately broad, glabrous (Fig. 119). Etymology. The species name is descriptive for the reticulated apical lobe of the style. Remarks. From D. spatulata Nielson, 1979 to which it is nearest to in shape of the style, D. reticulata sp.nov. can be distinguished by the much narrower caudodorsal process, absence of the caudoventral pygofer process and presence of reticulations on the pygofer caudodorsal process and apical stylar lobe.

Docalidia setosa sp.nov. (Figs 120–126, 151)

Type material. Holotype ♂: PERU: PA [Pasco]; Qbda., Castilla, 345 m., 10°10′S 75°15′W, 13.vi.87, P. Lozada (UNMSM). Description. Length. Male 8.85 mm, female unknown. External morphology. Large, robust species. General color light brown to black; forewings uniformly light brown; mesonotum black; pronotum dark brown; crown light

204 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia brown with dark markings; eyes light brown; face light to dark brown; clypeus dark brown with few small yellow spots; clypellus dark brown; lorae and genae light brown. Head short, wide, narrower than pronotum; anterior margin broadly rounded; crown short, broad, about as wide as width of eyes, slightly produced anteriorly, lateral margins nearly parallel, depressed on each side of middle; eyes large, semiglobular; clypeus long, broad, lateral margins broadly convex; clypellus broad, narrower than clypeus, lateral margins nearly parallel. Male genitalia. Male pygofer in lateral view large, glabrous, caudodorsal process moderately long, narrow, caudoventral process small, lobe-like (Fig. 120); segment X with short, narrow ventral process (Fig. 120); aedeagus very long, in lateral view shaft narrow, tubular, tapered to apex with long medial process on ventral margin, in dorsal view shaft inflated subbasally (Figs 121, 122); style long, robust, apical 1/5 large, profusely setaceous, apex pointed in dorsal view (Figs 123, 124) dorsal connective moderately long, narrow (Figs 121, 122); connective T-shaped, lateral arms narrow, horizontal, apices curved anteriorly, membrane and medial ridge absent, stem small, semiovate with narrow basal process on each side (Fig. 125); subgenital plate long, very broad, glabrous (Fig. 126). Etymology. The name of the species is descriptive for the large, profusely setose apex of the style. Remarks. Three species of Docalidia (D. spendlovei Nielson, 1979; D. zahniseri Nielson, 2011; D. parvitatis Nielson, 2011) have a similar style (large, setaceous apex) to D. setosa sp.nov. From them, D. setosa sp.nov. can be distinguished by the much narrower aedeagal shaft and far more dense microsetae on the style.

Docalidia trulla sp.nov. (Figs 127–133, 152)

Type material. Holotype ♂: PERU: MD [Madre de Dios], Z. R. [Zona Reservada], Tambopata, 290 m., 21.ii.87, P. Lozada (UNMSM). Description. Length. Male 8.50 mm, female unknown. External morphology. General color brown to black with numerous small, yellow markings. Forewing dark brown to black, veins with small yellow spots, cells with small, irregularly shaped, yellow markings; mesonotum black with very few small spots; pronotum black with numerous small, yellow bullae; crown ivory with few light brown markings; eyes dark brown; face light yellow with dark brown to black markings; clypeus with row of short, transverse black stripes on lateral margins, fused medially, few small, yellow spots; clypellus with suffused brown markings medially; lorae and genae with few, small black markings. Head short, broad, narrower than pronotum, broadly rounded anteriorly; crown broad, slightly narrower than eyes, produced anteriorly about 1/4 entire length beyond anterior margin of eyes, lateral margins convergent basally; eyes large, elongate ovoid; clypeus long, broadly convex; clypellus broad, narrower than clypeus, lateral margins slightly convex. Male genitalia. Pygofer in lateral view moderately large, subquadrate, glabrous, caudodorsal process long, narrow, caudoventral process small, lobe-like (Fig. 127);

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 205 M. W. NIELSON & P. W. LOZADA segment X with narrow ventral process (Fig. 127); aedeagus long, narrow, tubular, broadly curved in lateral view, ventral subapical process absent (Figs 128, 129); style extremely long, robust with narrow row of dense microsetae on lateral margin in distal 1/7, apex scoop-shaped (Figs 130, 131); dorsal connective short, narrow (Fig. 128); connective large, T-shaped, lateral arms narrow, without membrane and medial ridge, stem large, T-shaped (Fig. 132); subgenital plate very long, nearly broad throughout, glabrous (Fig. 133). Etymology. The name of the species (a noun in apposition) is descriptive for the scoop- shaped apex of the style. Remarks. From a group of Docalidia species having a robust style, much longer than aedeagus (D. cornicula Nielson, 2011; D. gigantea Nielson, 2011; D. longula Nielson, 2011; D. rondoniensis Nielson, 2011), D. trulla sp.nov. can be separated by the very narrow, tubular aedeagus in dorsal and lateral views and absence of the ventral subapical process.

New records for Peru Docalidia dentatula (Metcalf, 1964)

Material examined. MD [Madre de Dios], Z. R. [Zona Reservada], Tambopata (UNMSM).

Distribution. Bolivia (NIELSON 1979, 2011). Peru (new record).

Docalidia zanolae Nielson, 1986 Material examined. UC [Ucalyali], Pucallpa (UNMSM).

Distribution. Brazil (NIELSON 1986, 2011). Peru (new record).

Acknowledgements We greatly appreciate the reviews of this paper by Paul Freytag, retired Emeritus Professor University of Kentucky, Lexington and Murray Fletcher, Orange Agriculture Institute, Orange, NSW, Australia and for their useful comments and suggestions that improved the content of the paper. We especially thank Igor Malennovský for the invitation to join this Festschrift in honor of Pavel Lauterer and J. L. Stehlik.

References

NIELSON M. W. 1979: A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). III. Tribe Teruliini. Pacific Insects Monograph 35: 1–329. NIELSON M. W. 1982a: New species of Brazilian leafhoppers in the genus Docalidia Cicadellidae: Coelidiinae, Teruliini). Entomography 1: 237–256. NIELSON M. W. 1982b: New species of leafhoppers of Docalidia from Peru (Cicadellidae: Coelidiinae: Teruliini). Entomography 1: 289–320.

206 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

NIELSON M. W. 1982c: Some additional new species of leafhoppers of Docalidia from South America (Cicadellidae: Coelidiinae, Teruliini). Entomography 1: 439–445. NIELSON M. W. 1986: New South American leafhoppers in the genus Docalidia with a key to 37 species (Cicadellidae: Coelidiinae: Teruliini). Great Basin Naturalist 46(4): 749–762. NIELSON M. W. 1990: New species of Docalidia from the Neotropical region (Homptera: Cicadellidae: Coelidiinae). Entomologica Scandinavica 21(2): 225–240. NIELSON M. W. 1992: New species of Neotropical leafhoppers in the tribe Teruliini with a revised key to some species (Homoptera: Cicadellidae: Coelidiinae). Entomologica Scandinavica 21(2): 297–305. NIELSON M. W. 1996: Two new Neotropical genera and new species of Docalidia in the tribe Teruliini (Cicadellidae: Coelidiinae). Transactions of the American Entomological Society 122(2–3): 135–145. NIELSON M. W. 2011: New species in the Neotropical genus Docalidia with a key to known species, notes on distribution, taxonomy and a synoptic catalogue of the genus (Hemiptera: Cicadellidae: Coelidiinae: Teruliini). Zootaxa 2952: 1–86.

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Figs 1–7. Docalidia acucifera sp.nov. 1 – male pygofer, lateral view; 2 – aedeagus and dorsal connective, dorsal view; 3 – aedeagus and dorsal connective, lateral view; 4 – style, dorsal view; 5 – style, lateral view; 6 – connective, dorsal view; 7 – subgenital plate, ventral view.

208 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Figs 8–14. Docalidia basispinosa sp.nov. 8 – male pygofer, lateral view; 9 – aedeagus and dorsal connective, dorsal view; 10 – aedeagus and dorsal connective, lateral view; 11 – style, dorsal view; 12 – style, lateral view; 13 – connective, dorsal view; 14 – subgenital plate, ventral view.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 209 M. W. NIELSON & P. W. LOZADA

Figs 15–21. Docalidia bilobata sp.nov. 15 – male pygofer, lateral view; 16 – aedeagus and dorsal connective, dorsal view; 17 – aedeagus and dorsal connective, lateral view; 18 – style, dorsal view; 19 – style, lateral view; 20 – connective, dorsal view; 21 – subgenital plate, ventral view.

210 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Figs 22–28. Docalidia collaris sp.nov. 22 – male pygofer, lateral view; 23 – aedeagus and dorsal connective, dorsal view; 24 – aedeagus and dorsal connective, lateral view; 25 – style dorsal view; 26 – style, lateral view; 27 – connective, dorsal view; 28 – subgenital plate, ventral view.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 211 M. W. NIELSON & P. W. LOZADA

Figs 29–35. Docalidia crassitudinis sp.nov. 29 – male pygofer, lateral view; 30 – aedeagus and dorsal connective, dorsal view; 31 – aedeagus and dorsal connective, lateral view; 32 – style, dorsal view; 33 – style, lateral view; 34 – connective, dorsal view; 35 – subgenital plate, ventral view.

212 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Figs 36–42. Docalidia hama sp.nov. 36 – male pygofer, lateral view: 37 – aedeagus and dorsal connective, dorsal view; 38 – aedeagus and dorsal connective, lateral view; 39 – style, dorsal view; 40 – style, lateral view; 41 – connective, dorsal view; 42 – subgenital plate, ventral view.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 213 M. W. NIELSON & P. W. LOZADA

Figs 43–49. Docalidia iuncta sp.nov. 43 – male pygofer, lateral view; 44 – aedeagus and dorsal connective, dorsal view; 45 – aedeagus and dorsal connective, lateral view; 46 – style, dorsal view; 47 – style, lateral view; 48 – connective, dorsal view; 49 – subgenital plate, ventral view.

214 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Figs 50–56. Docalidia lineolaris sp.nov. 50 – male pygofer, lateral view; 51 – aedeagus and dorsal connective, dorsal view; 52 – aedeagus and dorsal connective, lateral view; 53 – style, dorsal view; 54 – style, lateral view; 55 – connective dorsal view; 56 – subgenital plate, ventral view.

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Figs 57–63. Docalidia maldonadoi sp.nov. 57 – male pygofer, lateral view; 58 – aedeagus and dorsal connective, dorsal view; 59 – aedeagus and dorsal connective, lateral view; 60 – style, dorsal view; 61 – style, lateral view; 62 – connective, dorsal view; 63 – subgenital plate, ventral view.

216 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Figs 64–70. Docalidia mediospinata sp.nov. 64 – male pygofer, lateral view; 65 – aedeagus and dorsal connective, dorsal view; 66 – aedeagus and dorsal connective, lateral view; 67 – style, dorsal view; 68 – style, lateral view; 69 – connective, dorsal view; 70 – subgenital plate, ventral view.

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Figs 71–77. Docalidia minatura sp.nov. 71 – male pygofer, lateral view; 72 – aedeagus and dorsal connective, dorsal view; 73 – aedeagus and dorsal connective, lateral view; 74 – style, dorsal view; 75 – style, lateral view; 76 – connective, dorsal view; 77 – subgenital plate, ventral view.

218 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and records of Docalidia

Figs 78–84. Docalidia minima sp.nov. 78 – male pygofer, lateral view; 79 – aedeagus and dorsal connective, dorsal view; 80 – aedeagus and dorsal connective, lateral view; 81 – style, dorsal view; 82 – style, lateral view; 83 – connective, dorsal view; 84 – subgenital plate, ventral view.

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Figs 85–91. Docalidia pakitzaensis sp.nov. 85 – male pygofer, lateral view; 86 – aedeagus and dorsal connective, dorsal view; 87 – aedeagus and dorsal connective, lateral view; 88 – style, dorsal view; 89 – style, lateral view; 90 – connective, dorsal view; 91 – subgenital plate, ventral view.

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Figs 92–98. Docalidia parabistyla sp.nov. 92 – male pygofer, lateral view; 93 – aedeagus and dorsal connective, dorsal view; 94 – aedeagus and dorsal connective, lateral view; 95 – style, dorsal view; 96 – style, lateral view; 97 – connective, dorsal view; 98 – subgenital plate, ventral view.

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Figs 99–105. Docalidia parainsolita sp.nov. 99 – male pygofer, lateral view; 100 – aedeagus and dorsal connective, dorsal view; 101 – aedeagus and dorsal connective, lateral view; 102 – style, dorsal view; 103 – style, lateral view; 104 – connective, dorsal view; 105 – subgenital plate, ventral view.

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Figs 106–112. Docalidia pedalis sp.nov. 106 – male pygofer, lateral view; 107 – aedeagus and dorsal connective, dorsal view; 108 – aedeagus and dorsal connective, lateral view; 109 – style, dorsal view; 110 – style, lateral view; 111 – connective, dorsal view; 112 – subgenital plate, ventral view.

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Figs 113–119. Docalidia reticulata sp.nov. 113 – male pygofer, lateral view; 114 – aedeagus and dorsal connective, dorsal view; 115 – aedeagus and dorsal connective, lateral view; 116 – style, dorsal view; 117 – style, lateral view; 118 – connective, dorsal view; 119 – subgenital plate, ventral view.

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Figs 120–126. Docalidia setosa sp.nov. 120 – male pygofer, lateral view; 121 – aedeagus and dorsal connective, dorsal view; 122 – aedeagus and dorsal connective, lateral view; 123 – style, dorsal view; 124 – style, lateral view; 125 – connective, dorsal view; 126 – subgenital plate, ventral view.

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Figs 127–133. Docalidia trulla sp.nov. 127 – male pygofer, lateral view; 128 – aedeagus and dorsal connective, dorsal view; 129 – aedeagus and dorsal connective, lateral view; 130 – style, dorsal view; 131 – style, lateral view; 132 – connective, dorsal view; 133 – subgenital plate, ventral view.

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Figs 134–142. Docalidia spp., general habitus. 134 – D. acucifera sp.nov.; 135 – D. basispinosa sp.nov.; 136 – D. bilobata sp.nov.; 137 – D. collaris sp.nov.; 138 – D. crassitudinis sp.nov.; 139 – D. hama sp.nov.; 140 – D. iuncta sp.nov.; 141 – D. lineolaris sp.nov.; 142 – D. maldonadoi sp.nov.

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Figs 143–152. Docalidia spp., general habitus. 143 – D. mediospinata sp.nov.; 144 – D. minatura sp.nov.; 145 – D. minima sp.nov.; 146 – D. pakitzaensis sp.nov.; 147 – D. parabistyla sp.nov.; 148 – D. parainsolita sp.nov.; 149 – D. pedalis sp.nov.; 150 – D. reticulata sp.nov.; 151 – D. setosa sp.nov.; 152 – D. trulla sp.nov.

228 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 229–234, 2013

Platymetopius pavelitus sp.nov. (Hemiptera: Auchenorrhyncha: Cicadellidae), a new species from Italy

VERA D’URSO1, CHRISTOPH BÜCKLE2 & ADALGISA GUGLIELMINO3 1 Department of Biological, Geological and Enviromental Sciences, sez. Animal Biology, University of Catania, via Androne 81, I-95124 Catania, Italy; e-mail: [email protected] 2 Neckarhalde 48, D-72070 Tübingen, Germany

3 Department of Agriculture, Forests, Nature and Energy, University of Tuscia, Viterbo, 01100 Italy; e-mail: [email protected]

D’URSO V., BÜCKLE C. & GUGLIELMINO A. 2013: Platymetopius pavelitus sp.nov. (Hemiptera: Auchenorrhyncha: Cicadellidae), a new species from Italy. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 229–234. – Platymetopius pavelitus sp.nov. is described from southern Italy (Calabria and Sicily). Its diagnostic characters lie in the structure of the male genitalia (aedeagus, pygofer appendages) and in the female pregenital sternite. Keywords. Cicadomorpha, , taxonomy, Palaearctic Region, Sicily, Calabria

Introduction The genus Platymetopius Burmeister, 1838 was originally established for the species Cicada vittata Fabricius, 1775 (= Platymetopius undatus De Geer, 1773, type species) and Jassus rostratus Herrich-Schäffer, 1834 (BURMEISTER 1838). Currently it includes more than 75 species distributed in the Palaearctic and Oriental regions. However, the genus as currently defined is probably not monophyletic. It is one of the largest and most complex genera of the Deltocephalinae and urgently in need of revision. At present, it is divided in two subgenera: Platymetopius s. str. (73 species) and Quernus Dlabola, 1974 (3 species) (ZAHNISER 2013). ABDUL-NOUR (2001) established five informal species-groups among the Platymetopius taxa present in the Near East, based on morphological characters (e.g. body shape, coloration, aedeagus morphology, structure of pygofer appendages and female pregenital sternite). In 1977, R. Remane collected one male and two females of an unknown Platymetopius species on Mount Etna (Sicily). Subsequently, in 1985, one of the authors (V. D’Urso) found one male in the Peloritani Mountains (Sicily) and, in 2009, the other two authors (A. Guglielmino and Ch. Bückle) collected three females in the Sila Mountains (Calabria), all belonging to the same taxon. Based on external morphology and the structures of the male genitalia and the female pregenital sternite, these specimens are considered as belonging to a new species, here described as Platymetopius pavelitus sp.nov.

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Material and methods The specimens examined are deposited in the institutions abbreviated as: ABUC ...... Department of Biological, Geological and Environmental Sciences, sez. Animal Biology, University of Catania, Italy (coll. V. D’Urso) AFUT ...... Department of Agriculture, Forests, Nature and Energy, University of Tuscia, Viterbo, Italy (coll. A. Guglielmino) SMTD ...... Senckenberg Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Dresden, Germany (coll. R. Remane)

The morphological terminology used in the description follows OSSIANNILSSON (1978). Measurements were taken with the aid of a micrometer attached to a Zeiss SV8 stereomicroscope.

Taxonomy Platymetopius pavelitus sp.nov. (Figs 1–9)

Type material. Holotype: ♂, Italy, Sicily, Etna, supra Linguaglossa (CT), 37°48′51″N 15°05′43″E, 1100 m, 21.09.1977, Leguminosae (Cytisus?), , Quercus cerris, Q. pubescens, Farn [fern], Weißdorn (lockerwüchsig) [ (sparse)], R. Remane leg. et coll. (SMTD). Paratypes: 2 ♀♀, same data as holotype (SMTD); 1 ♂, Italy, Sicily, Mt. Peloritani, Portella Fossa Lupo (Mandanici-ME), 885 m, 1.10.1985, clearing in Quercus forest, D’Urso leg. et coll. (ABUC); 3 ♀♀, Italy, Calabria, Sila, SS 660, km 35.5 (Acri-CS), 1150 m, 13.08.2009, open area near Pinus afforestation, Guglielmino & Bückle leg., coll. A. Guglielmino (AFUT). Description. Measurements. Males. Holotype. Total body length (including tegmina): 4.90 mm; vertex length: 0.32 mm; head width: 1.40 mm; pronotum length: 0.65 mm; pronotum width: 1.43 mm; length of vertex + pronotum + mesonotum: 1.60 mm. Paratype. Total body length (including tegmina): 5.06 mm; vertex length: 0.32 mm; head width: 1.44 mm; pronotum length: 0.68 mm; pronotum width: 1.44 mm; length of vertex + pronotum + mesonotum: 1.60 mm. Females. Paratypes. Total body length (including tegmina): 5.72–5.93 mm; vertex length: 0.36–0.38 mm; head width: 1.52–1.65 mm; pronotum length: 0.72–0.82 mm; pronotum width: 1.64–1.80 mm; length of vertex + pronotum + mesonotum: 1.60–1.80 mm. All specimens examined are macropterous with the forewings longer than the abdomen and as long as the hind wings (Figs 1, 2). General coloration of body is yellowish-brown, mottled with dark brownish dots and with hyaline spots on the forewings; the dark pattern is paler in the females (Fig. 2). Head as wide as pronotum (male) or slightly wider (females). Vertex with rounded, slightly elevated anterior margin and a longitudinal median ridge in the posterior third; its width between the eyes twice the median length; four brown spots along the anterior margin, one near each ocellus. Frons slightly convex. Face yellowish, in males with light brown transverse stripes on frons. Gena with one small, brown spot near ocellus, usually with a small, dark brownish dot near lorum. Pronotum more than twice wider than long, mottled with brown dots. Scutellum yellowish, with two brown spots anterior to the brown cicatrix. Forewings densely mottled with brown dots and with large hyaline areas.

230 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Platymetopius pavelitus sp.nov.

Figs 1–2. Platymetopius pavelitus sp.nov., paratypes, habitus. 1 – male (from Peloritani Mts.); 2 –female (from Sila).

Male genitalia. Pygofer (Fig. 3) with a deep semicircular notch on lateral portion of posterior margin of its lobes; notch in its dorsal portion with a short spine-shaped process. Each lobe with a group of about 15 long and short setae. Anal tube as in Fig. 3. Genital valve triangular (Fig. 4), less than twice as wide as long, with anterior margin slightly concave in its median portion, lateral margin slightly convex in its median portion. Genital plate as in Fig. 4, with median margin shorter than length of the genital valve and lateral margin convex in its basal half. Aedeagus slender with a pair of lateral appendages arising from the basal portion of the shaft (Figs 3, 5, 6); shaft and appendages slender, curved dorsad; appendages longer than shaft;) aedeagus base anchor-shaped in posterior view (Fig. 6). Connective and styles as in Figs 7 and 8. Female genitalia. Hind margin of pregenital sternite deeply concave with a median bifurcate process (Fig. 9). Etymology. It is our pleasure to name the species in honour of Pavel Lauterer, a famous and excellent researcher of Hemiptera and a good friend of ours. The name “pavelitus” should be treated as a noun in apposition.

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Differential diagnosis. Platymetopius pavelitus sp.nov. differs from the other species of the genus particularly in the morphology of the aedeagus, the pygofer appendages and the female pregenital sternite (see also Remarks). Biological notes. The specimens were collected in summer and early autumn in low vegetation near pine (Pinus) and oak (Quercus) woods. Distribution. Up to now, P. pavelitus sp.nov. has been collected only in Calabria and Sicily. Remarks. P. pavelitus sp.nov. is characterized by the following characters: yellowish- brown coloration mottled with dark dots and with hyaline areas on the forewings; short and rounded vertex with four dark spots; pygofer lobes with semicircular notch and short spine-shaped process; slender aedeagus with one pair of long, lateral appendages arising from the base of the shaft; posterior margin of the female pregenital sternite deeply concave with median bifurcate process. On the basis of those characters, the new species may be placed within the subgenus Platymetopius s. str. P. pavelitus sp.nov. resembles P. exhereditus Dlabola et Heller, 1962, known from Iran, in the shape and coloration of the vertex, the short pygofer appendages and the slender aedeagus with only one pair of long appendages. It differs from that species in the antero-ventral direction of the pygofer process (directed dorso-caudad in P. exhereditus), the semicircular notch below the pygofer process (pygofer margin is convex in P. exhereditus), the subapical position of the pygofer process (apical in P. exhereditus) and the shorter aedeagus appendages not covering the shaft in lateral view (longer than the shaft and covering it for most of its length in P. exhereditus). The new species shares some characters with the Platymetopius obsoletus group as defined by ABDUL-NOUR (2001). In particular, P. pavelitus resembles P. obsoletus (Signoret, 1880) sensu Haupt, 1927 and other closely-related species (P. ferrarii Haupt, 1927, P. cruentatus Haupt, 1927 and P. retamae Linnavuori, 1972) in the general habitus (forewing coloration without zig-zag pattern, vertex rounded and quite short with dark spots along its anterior margin) and in the presence of only one pair of aedeagus appendages. However, P. pavelitus sp.nov. differs in the shape of the pygofer appendages (slender and long in the species of the obsoletus group sensu ABDUL-NOUR (2001), except for P. retamae which, however, presents a different shape of pygofer appendages with respect to the new species), in the slender base of the aedeagus and in the female pregenital sternite with a median bifurcate posterior process (trilobate posterior margin in the species of obsoletus group).

Acknowledgements We are grateful to Igor Malenovský (Moravian Museum, Brno, Czech Republic) and James Zahniser (Illinois Natural History Survey, Champaign, Illinois, USA) for their invaluable suggestions on the manuscript and to Christian Schmidt (Senckenberg Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Dresden, Germany) for the loan of specimens from the R. Remane collection.

232 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Platymetopius pavelitus sp.nov.

Figs 3–9. Platymetopius pavelitus sp.nov. 3–8 – Paratype, male (from Peloritani Mts.): 3 – genital block, lateral view; 4 – genital plates and genital valve, ventral view; 5 – aedeagus, right lateral view; 6 – aedeagus, posterior view; 7 – connective, dorsal view; 8 – left stylus, dorsal view. 9 – Paratype, female (from Sila), pregenital sternite, ventral view.

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References

ABDUL-NOUR H. 2001: Le genre Platymetopius Burmeister, 1838 au Proche-Orient: inventaire et descriptions d’espèces nouvelles ou peu connues (Hemiptera, Cicadomorpha, Cicadellidae). Nouvelle Revue d’Entomologie (N.S.) 18(1): 77–89. BURMEISTER H. 1838: Genera insectorum iconibus illustravit et descripsit. Volumen I. Rhynchota. Burmeister et Stange, Berolini, 76 pp., pls. 10, 11, 17, 20. DLABOLA J. 1974: Ergebnisse der tschechoslowakisch-iranischen entomologischen Expedition nach dem Iran im 1970. N3. Homoptera Auchenorrhyncha (1. Teil). Acta Entomologica Musei Nationalis Pragae Suppl. 6: 29–73. OSSIANNILSSON F. 1978: The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark. Part I. Fauna Entomologica Scandinavica 7(1): 1–222. ZAHNISER J. N. 2013: 3i interactive keys and taxonomic databases, subfamily Deltocephalinae. Available online: http://imperialis.inhs.illinois.edu/zahniser/ (accessed on 9 August 2013).

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New records of Auchenorrhyncha (Hemiptera) for the Czech Republic

IGOR MALENOVSKÝ Department of Entomology, Moravian Museum, Hviezdoslavova 29a, CZ-627 00 Brno, Czech Republic; e-mail: [email protected], [email protected]

MALENOVSKÝ I. 2013: New records of Auchenorrhyncha (Hemiptera) for the Czech Republic. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 235–263. – Faunistic data on 17 species of planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) from the Czech Republic are provided and discussed. Five species are recorded from the Czech Republic for the first time: Javesella bottnica Huldén, 1974; Litemixia pulchripennis Asche, 1980; Cicadella lasiocarpae Ossiannilsson, 1981; ohausi Wagner, 1939; and coronifer (Marshall, 1866). Diagnostic characters of Javesella bottnica, a boreal species which is recorded for the first time in central Europe, are briefly discussed and the species is compared with Javesella simillima (Linnavuori, 1948); habitus of male and female brachypterous specimens and the male anal tube and the aedeagus are illustrated for both species. Seven species (previously known in the Czech Republic only from Moravia) are newly recorded for Bohemia: Kelisia confusa Linnavuori, 1957; Kelisia guttulifera (Kirschbaum, 1868); Eupteryx lelievrei (Lethierry, 1874); Zyginidia pullula (Boheman, 1845); Allygidius abbreviatus (Lethierry, 1878); septentrionalis Wagner, 1963; and Pinumius areatus (Stål, 1858). Four species (previously known in the Czech Republic only from Bohemia) are newly ascertained also for Moravia: Anoscopus alpinus (Wagner, 1955); Chlorita dumosa (Ribaut, 1933); Eupteryx signatipennis (Boheman, 1847); and Jassargus allobrogicus (Ribaut, 1936). Records of an additional little-known species in the Czech Republic, Hephathus achilleae Mityaev, 1967, are also provided. Keywords. Fulgoromorpha, Cicadomorpha, Delphacidae, Cicadellidae, Planthoppers, leafhoppers, Javesella, distribution, taxonomy, central Europe, Czech Republic, Bohemia, Moravia, France

Introduction The earliest beginnings of studies on the Czech fauna of the Auchenorrhyncha date back to the end of the 18th century. After pioneering contributions had been made by polymath naturalists and early entomologists (PREYSSLER 1792, PREYSSLER et al. 1793; KOLENATI 1859, 1860), three leading Auchenorrhyncha taxonomists, Franz Xaver Fieber (1807–1872), Leopold Melichar (1856–1924), and Franz Then (1841–1919), in part, were active on the territory of what is today the Czech Republic (EMMRICH 2003, BEZDÌK 2011). The first comprehensive lists of species that were known to occur in the country were provided by DUDA (1892) and SPITZNER (1892) for both large historical parts of the Czech Lands, Bohemia and Moravia, respectively. The pace of knowledge acquisition on the presence and distribution of Auchenorrhyncha species picked up sharply after the 1940s, through the activities and numerous publications of Veleslav Lang (1913–1993), Jiøí Dlabola (*1922) and Pavel Lauterer (*1933) (see JEŽEK 1994, KOLEŠKA 1998, MALENOVSKÝ & BURCKHARDT 2003 and MALENOVSKÝ 2013 for biographies and bibliographies). The rapidly burgeoning quantities of data were summed up in a monograph (DLABOLA 1954) and a check-list (DLABOLA 1977) of the Czechoslovak

235 I. MALENOVSKÝ fauna, both of them major milestones. Continuous field work, identification and revisions of museum collections, improving knowledge of the taxonomy and biology of many central European species and the expansion of exotic species, have lead to discoveries of additional Auchenorrhyncha species in recent years as well (e.g. MALENOVSKÝ 2006; MALENOVSKÝ & LAUTERER 2006, 2010, 2012; MALENOVSKÝ & TROPEK 2009). This paper follows MALENOVSKÝ & LAUTERER (2010) in form and its aim of reporting data on the occurrence of some hitherto insufficiently-known Auchenorrhyncha species in the Czech Republic, including some first records, as one step in the preparation of a new, up-to-date Hemiptera check-list for the country. To date, some 580 species of Auchenorrhyncha (including those in the present paper) have been reported for the Czech Republic, partly scattered in the literature; revisions of some critical taxa and old doubtful records are, however, still required, and records of some additional species, especially those already known from neighbouring countries but not from the Czech Republic, may still be anticipated.

Material and methods The material examined or cited in the paper is deposited in the following institutions, abbreviated as: BMNH ...... Natural History Museum, London, United Kingdom MMBC ...... Moravian Museum, Brno, Czech Republic MNHN ...... Muséum national d’Histoire naturelle, Paris, France NMPC ...... National Museum, Prague, Czech Republic SWHG ...... private collection of Sabine Walter, Kurort Hartha, Germany.

For the purposes of faunistic research, the territory of the Czech Republic is traditionally divided into two parts: Bohemia in the west and Moravia (including former Austrian, later Czech, Silesia) in the east (e.g. DLABOLA 1977, BOGUSCH et al. 2007, KMENT 2009). These parts largely correspond to the politico-historical arrangement of the Czech Lands in the Middle Ages through to the times of the Habsburg monarchy, as well as of before 1949. They are not, however, reflected in the current administrative status of the Czech Republic. The boundary between Bohemia and Moravia is thus interpreted here in the historical sense, precisely in the form established in 1924 (see KMENT 2009 for details). The localities cited in the “Material examined” sections are ordered according to their field code numbers (given in parentheses after the names of conurbations) in the faunistic and floristic grid mapping system of central Europe (EHRENDORFER & HAMANN 1965, PRUNER & MÍKA 1996). The same mapping system is also used in Figs 9–20.

The following abbreviations are also used to refer to localities and material examined: NM – Nature Monument, NNM – National Nature Monument, NNR – National Nature Reserve, NR – Nature Reserve (all of these are the various categories contained within the Czech system of legislatively protected small-scale natural areas); det. – determined by, leg. – collected by.

236 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

All material was identified by I. Malenovský unless stated otherwise. The nomenclature and classification follow HOCH (2013) and TISHECHKIN (1999). Photographs of specimens were taken using a Leica DFC 295 digital camera on a Leica Z16 APO macroscope and the Leica Application Suite software or an Olympus 5060WZ camera on an Olympus BX41 microscope and the QuickPhoto Camera software. Eye- piece graticule on a MBS 10 stereomicroscope was applied for measurements of dry- mounted specimens.

Results FULGOROMORPHA

Delphacidae

Javesella bottnica Huldén, 1974 (Figs 1, 2, 5, 7, 9) Material examined. Bohemia: Pøebuz, 2.8 km NW, Rolavská vrchovištì NNR, Lieche (=Rolavský rybník Pond) (5641), 50°23′32″N, 12°37′41″E, 910 m a.s.l., 21.vi.2011, peatbog, 3 ♂♂ 7 ♀♀ (all brachypterous), I. Malenovský leg. (MMBC). Remarks. A boreal species so far known only from Finland and northern Russia (Karelia) (HULDÉN 1974, OSSIANNILSSON 1978, SÖDERMAN 2007) where it was swept from sedge (Carex acuta) bordering an oligotrophic lake, and from lakeshore (SÖDERMAN 2007). The morphology of the male genitalia is similar to Javesella simillima (Linnavuori, 1948) and J. discolor (Boheman, 1847). HOLZINGER et al. (2003) considered J. bottnica and J. simillima as possibly conspecific. Besides differences in the shape of the male genitalia (not specified explicitly), HULDÉN (1974) distinguished J. bottnica from J. simillima and J. discolor also based on more elongated wings in brachypterous specimens (ratio wing length/width in males 1.79–2.32 in J. bottnica, 1.57–1.68 in J. simillima, and 1.43–1.82 in J. discolor). It also comes out from the original description that J. bottnica is larger than J. simillima (body length of brachypterous specimens: J. bottnica – 2.00–2.46 mm (males), 2.66 mm (female); J. simillima (cited after LINNAVUORI 1969) – 1.5–1.6 mm (males), 2.0–2.1 mm (females)). A series of brachypterous specimens collected recently in north-western Bohemia, on the Czech side of the Krušné hory Mts. [Erzgebirge], at the Lieche Pond near the town of Pøebuz (Fig. 9), in a complex of peat- and intermediary bogs and littoral vegetation (dominated by Eriophorum angustifolium, E. vaginatum, Carex limosa, C. rostrata and Molinia caerulea; Fig. 7) perfectly fits to the original description by HULDÉN (1974) and illustrations of J. bottnica by OSSIANNILSSON (1978; reprinted also in HOLZINGER et al. 2003). A comparison with a series of J. simillima collected in adjacent part of Germany (Saxony: Bad Brambach, Gründel, 3.vii.2004, 4 ♂♂ 4 ♀♀; Bad Brambach, Raunerbachtal/Gründel, 16. and 18.vi.2005, 2 ♂♂ 3 ♀♀, Pern, Hammerlug, 9.vi.2011; 5 ♂♂ 6 ♀♀; all brachypterous specimens, S. Walter leg. et det., SWHG) confirmed this identification and the differences between both species outlined by previous authors (see also SÖDERMAN 2007). The specimens of J. bottnica from Pøebuz: Lieche (n = 3 ♂♂,

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Figs 1–6. Javesella spp.: 1, 2, 5 – J. bottnica Huldén, 1974 (specimens from the Czech Republic: Krušné hory Mts., Pøebuz, Lieche Pond); 3, 4, 6 – J. simillima (Linnavuori, 1948) (specimens from Germany: Saxony, Bad Brambach, Gründel). 1, 3 – brachypterous males, habitus, dorsal view; 2, 4 – brachypterous females, habitus, dorsal view; 5, 6 – male anal tube and aedeagus, left lateral view.

238 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Figs 7–8. Habitats of some planthopper species recorded in the Czech Republic for the first time. 7 – complex of intermediary and peat-bogs at the Lieche Pond, Rolavská vrchovištì National Nature Reserve near Pøebuz in the Krušné hory Mts., north-western Bohemia: the locality of Javesella bottnica Huldén, 1974 (photograph by J. Sychra); 8 – calcareous spring fen at the Olšový rybník Pond near Louèeò in central Bohemia: one of the Czech localities of Litemixia pulchripennis Asche, 1980 and Kelisia confusa Linnavuori, 1957 (photograph taken on a field course in entomology by L. Hubáèková). Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 239 I. MALENOVSKÝ

7 ♀♀) are generally larger (body length: males: 2.48–2.58 mm; females: 2.55–2.85 mm) and have relatively longer wings (ratio wing length/width: males: 1.81–1.97; females: 1.91–2.09) than specimens of J. simillima from Saxony (n = 8 ♂♂, 11 ♀♀; body length: males 1.65–1.85 mm; females: 2.00–2.25 mm; ratio wing length/width: males: 1.46–1.64; females: 1.37–1.77); the body, particularly the head and thorax, of both sexes of J. bottnica are also slightly darker than the examined material of J. simillima (Figs 1–4). Both species also differ in the shape of the appendages of the male anal tube in lateral view (nearly straight in J. bottnica, curved in J. simillima) and the shape of the aedeagus in lateral view (with a narrow apex, sinuate posterior margin subapically and sharp tooth on anterior margin in J. bottnica, and a broad apex, nearly straight posterior margin subapically and an obtuse tooth on anterior margin in J. simillima) as illustrated here in Figs 5–6. As these differences appear constant across a large geographical distance (Fennoscandia and central Europe), J. bottnica is considered here as a valid species distinct from J. simillima and also J. discolor which, besides the shorter wings of brachypterous specimens, differs from J. bottnica by strongly curved appendages of the male anal tube and a broader apex of the aedeagus in lateral view (illustrated in OSSIANNILSSON 1978: p. 195 and HOLZINGER et al. 2003: p. 310). Several specimens of J. discolor matching the description by OSSIANNILSSON (1978) were recorded in the Krušné hory Mts. on the same site as the specimens of J. bottnica but in a different habitat (a Nardus stricta-Avenella flexuosa grassland at a spruce forest margin). More field work is needed to elucidate the biology of J. bottnica. Based on the data in literature and the present finding, the species is probably associated with Carex and/or Eriophorum spp. in cold and wet, bog habitats. Javesella bottnica is recorded here for the first time for the Czech Republic and central Europe. Its occurrence in this region is probably relict, perhaps from glacial times, as in many other tyrphobiont or tyrphophilous insects (e.g. SPITZER & DANKS 2006).

Kelisia confusa Linnavuori, 1957 (Figs 8, 9, 21)

Published records. MALENOVSKÝ & LAUTERER (2010): Bedøichov, towards Èernovice (6564); Brno-Jehnice, Ponávka Brook valley (6765); Brno-Chrlice, Splavisko (6865); Rašovice, Rašovický zlom-Chobot NR (6867), Dolní Dunajovice, River Dyje floodplain (7165); Knìždub, Èertoryje NNR (7170); Radìjov, Kútky NR (7170); Hlohovec, Allahovy rybníky Ponds (7266); Sedlec, Slanisko u Nesytu NNR (7266). Material examined. Bohemia: Louèeò, 1 km NW, shore and alder carr at Knížecí rybník Pond (5756), 50°17′30″N, 15°00′29″E, 253 m a.s.l., 21.viii.2012, 3 ♀♀. Chudíø, 1 km SE, spring fen and shore of Olšový rybník Pond (5756), 50°18′06″N, 15°01′04″E, 235 m a.s.l., 21.viii.2012, 1 ♂ 1 ♀. All specimens I. Malenovský leg. (MMBC). Remarks. In tall-sedge swamps near the shores of ponds or the floodplains of streams, in fens and along bog margins at moderately eutrophic sites, probably associated with Carex elata and/or C. acutiformis (HOLZINGER et al. 2003, NICKEL 2003, MALENOVSKÝ & LAUTERER 2010). Known from central Europe, Lithuania, Finland, Italy, and the Balkan peninsula (HOLZINGER et al. 2003, SÖDERMAN 2007, HOCH 2013). In the Czech Republic, documented previously only from southern Moravia (MALENOVSKÝ & LAUTERER 2010). Recorded here for the first time for Bohemia: the River Elbe lowlands in central Bohemia (Fig. 9).

240 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Fig. 9. Distribution of Javesella bottnica Huldén, 1974 (cross) and Kelisia confusa Linnavuori, 1957 (circles) in the Czech Republic.

Kelisia guttulifera (Kirschbaum, 1868) (Fig. 10)

Published records. DLABOLA (1956): Hostýn (6672). MALENOVSKÝ & LAUTERER (2012): Nedašov, Pod Cigánem NM (6974); Starý Hrozenkov, Hrozenkovské terasy (7073). Material examined. Bohemia: Louèeò, 1 km NW, alder carr wood on shore of Knížecí rybník Pond (5756), 50°17′30″N, 15°00′29″E, 253 m a.s.l., 21.viii.2012, on Carex remota,1 ♂, I. Malenovský leg. (MMBC). Moravia: Hostýn (6672), vii.1954, 1 ♂, Starý leg., J. Dlabola det., I. Malenovský revid. (MNHN). Babice nad Svitavou, Myší díra and Svitava River valley (6765), 240–300 m a.s.l., 2.x.1999, 1 ♀, P. Lauterer leg. (MMBC). Remarks. In moist places in oak and beech forests (often on or along abandoned tracks) and in alder fen woodland on Carex sylvatica, C. remota and C. elongata as well as in open wetlands such as spring mires, wet meadows and sandpits on C. vulpina, C. distans and C. paniculata (HOLZINGER et al. 2003, NICKEL 2003). Widespread in western, central and southern Europe from Great Britain to Lithuania in the north and from Corsica and Sardinia to the Balkan peninsula in the south (HOLZINGER et al. 2003, HOCH 2013). Poorly documented in the Czech Republic: overlooked or occurring only locally. Previously known from only three records from springs fens in the Hostýnské vrchy Hills and the Bílé Karpaty Mts. in eastern Moravia (DLABOLA 1956, MALENOVSKÝ & LAUTERER 2012). New species for Bohemia (Fig. 10).

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Fig. 10. Distribution of Kelisia guttulifera (Kirschbaum, 1868) in the Czech Republic.

Litemixia pulchripennis Asche, 1980 (Figs 8, 11, 22) Material examined. Bohemia: Lysá nad Labem, 2 km N, Hrabanovská èernava NNM (5755), 50°13′02″N; 14°50′09″E, 185 m a.s.l., 23.viii.2012, a large open lowland fen, 5 ♂♂ 4 ♀♀. Louèeò, 1 km NW, calcareous spring fen upstream Knížecí rybník Pond (5756), 50°17′26″N, 15°00′19″E, 254 m a.s.l., 21.viii.2012, 7 ♂♂ 17 ♀♀. Chudíø, 1 km SE, calcareous spring fen at Olšový rybník Pond (5756), 50°18′06″N, 15°01′04″E, 235 m a.s.l., 21.viii.2012, 6 ♂♂ 4 ♀♀. All specimens I. Malenovský leg. (MMBC). Remarks. A rarely-documented species with just a handful of records from south- western, central and north-eastern France (ASCHE 1980, REMANE & DELLA GIUSTINA 1991, DELLA GIUSTINA & REMANE 1992); there is one more unpublished record from the northern foothills of the Vosges Mts., Moselle (57): Philippsbourg, Etang de Hanau, 49°00′37″N, 7°32′16″E, 234 m a.s.l., 16.viii.1995, peat-bog on shore of a pond, several ♂♂ ♀♀, I. Malenovský leg., MMBC), southern Germany (northern upper Rhine plain and Bavaria, mostly in open pine forests on temporarily moist, acidic and sandy substrates in lowlands: REMANE & FRÖHLICH 1994, NICKEL 2003, KUNZ et al. 2011), and a single record in Austria in an intermittently wet meadow in the environs of Vienna (HOLZINGER 2009a, b). All three Czech records were made in relict open (treeless) calcareous fen habitats in the River Elbe lowlands in central Bohemia (Figs 8, 11), where the species was invariably collected from Molinia caerulea (sensu stricto, DANÈÁK et al. 2012). Molinia caerulea is the single confirmed host plant (REMANE & DELLA GIUSTINA

242 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Fig. 11. Distribution of Litemixia pulchripennis Asche, 1980 (crosses), Cicadella lasiocarpae Ossiannilsson, 1981 (empty circle), Chlorita dumosa (Ribaut, 1933) (squares), Euscelis ohausi Wagner, 1939 (full circle), and Pinumius areatus (Stål, 1858) (diamonds) in the Czech Republic.

1991, NICKEL 2003); even if some published records associate other taxa of this taxonomically difficult aggregate of grasses, the upper Rhine plain populations in Germany, at the least, were collected from M. caerulea (H. Nickel pers. comm.). As adults of L. pulchripennis usually remain deep in the tussocks of the host plant, only very few specimens can be taken with a sweep-net; most of the material cited above was collected using a petrol-driven suction sampler (G-Vac; STEWART 2002) which is probably the most efficient method for monitoring L. pulchripennis. Most specimens collected are brachypterous, which suggests permanent populations at the sites. New species for the Czech Republic.

CICADOMORPHA

Cicadellidae: Macropsinae

Hephathus achilleae Mityaev, 1967 (Figs 12, 23)

Published data. TISHECHKIN (1999): Southern Moravia. Material examined. Moravia: Mohelno (6863), 25.vii.1942, 2 ♂♂ and 20.vii.1943, 1 ♂, both J. Šnoflák leg. Brno-Slatina, Stránská skála hill (6866), 22.vii.1954, dry grassland on limestone, 7 ♂♂ 2 ♀♀, P. Lauterer leg.

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Køižanovice (6867), 10.vii.1963, 4 ♂♂ 1 ♀, L. Pospíšilová leg. Miroslav (7063), 14.vii.1963, 2 ♀♀, L. Pospíšilová leg. Trstìnice near Znojmo, grassland above the village (7063), 6.viii.1965, 2 ♀♀, L. Pospíšilová leg. Popice (7065), 16.vi.1977, 2 ♂♂, M. Kocourek leg. Pouzdøany (7065), 2.vii.1977, 1 ♂, M. Kocourek leg. Èejè (7067), vi.1940, 22 ♂♂ 2 ♀♀ (MNHN), vii.–viii.1940, 4 ♂♂ (NMPC), all A. Hoffer leg. Kobylí (7067), vii.1975, 3 ♂♂, M. Kocourek leg.; 183–274 m a.s.l., 19.vii.1977, 1 ♂, dry grassland, L. Pospíšilová leg. Sedlec, Kamenný vrch Hill [= Skalky] (7266), 25.vii.1968, 2 ♂♂, P. Lauterer leg. All material in MMBC, if not indicated otherwise.

Remarks. Described from Kazakhstan and recently redescribed by TISHECHKIN (1999), who also recorded it from central and southern parts of European Russia, northern Caucasus, Georgia, and central Europe: Hungary and the Czech Republic (southern Moravia). Details of the Czech localities were not given in the paper; the information was based on specimens from Kobylí (7067), deposited in BMNH (D. Tishechkin, pers. comm.). Later, the species was also confirmed for eastern Finland (SÖDERMAN 2007). H. achilleae is reported as living on steppes, dry meadows and dry river banks, mainly upon various Asteraceae: Artemisia campestris, A. absinthum, A. albicerata, A. santolinifolia, Erigeron acris, Achillea millefolium, Tanacetum sp. and, perhaps accidentally, on Spiraea hypericifolia (Rosaceae) (TISHECHKIN 1999, SÖDERMAN 2007). All known Moravian localities are well-preserved dry grassland sites on the northern outskirts of the Pannonian lowlands, or close to them (Fig. 12). Hephathus achilleae is probably more widespread in central and eastern Europe (there are additional unpublished data in the collections of MMBC, MNHN and NMPC from Slovakia, at the least) but may have been partly confused with the similar and closely related H. nanus (Herrich-Schäffer, 1835); the two species can be differentiated mainly by the colour pattern of the face (as especially darker forms may be difficult to identify, it is helpful to have a series of specimens from a given locality) and acoustic signals (TISHECHKIN 1999). However, all specimens of H. achilleae cited here from southern Moravia were collected more than 35 years ago and H. achilleae has not been rediscovered despite relatively extensive faunistic surveys of dry grassland sites in southern Moravia in recent years. The species may thus have declined or become extinct in the Czech Republic, in contrast to H, nanus which has recently been confirmed from a number of localities (MALENOVSKÝ et al. 2011, MALENOVSKÝ & LAUTERER 2012).

Cicadellidae:

Anoscopus alpinus (Wagner, 1955) (Figs 13, 24)

Published data. LAUTERER & NOVOTNÝ (1991): Krkonoše Mts., Mt Snìžka (5260). Material examined. Bohemia: Pøebuz, Pøebuzské vøesovištì NM (5641), 50°22′38″N, 12°36′43″E, 880 m a.s.l., 22.vi.2011, secondary montane heath on former peat diggings, 2 ♂♂. Moravia: Staré Mìsto, 9 km NW, summit of Mt Králický Snìžník (5767), 50°12′27″N, 16°50′51″E, 1400–1420 m a.s.l., 7.vii.2006, alpine grassland, under tussocks of Festuca supina, 2 ♂♂ 4 ♀♀. Karlova Studánka, 6 km SW, Pradìd NNR, Mt Vysoká hole (5969), 50°03′41″N, 17°14′15″E, 1450 m a.s.l., 17.viii.2011, alpine grassland, under tussocks of Festuca supina, Nardus stricta and Avenella flexuosa,4 ♀♀ 2 nymphs. All material I. Malenovský leg. (MMBC). Remarks. Known from the mountains of Austria, Germany, Switzerland, Poland, and the Czech Republic but perhaps conspecific with the western Mediterranean A. assimilis

244 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Fig. 12. Distribution of Hephathus achilleae Mityaev, 1967 in the Czech Republic.

(Signoret, 1879) and A. duffieldi Le Quesne, 1964 described from Great Britain (see e.g. REMANE & FRÖHLICH 1994 for a discussion of taxonomy). Probably lives on grasses in heaths and bogs from the submontane to the alpine belt (NICKEL 2003). The series from the Králický Snìžník Mts. was collected by hand from under the tussocks of an almost monodominant growth of Festuca supina, while adults in Pøebuz were swept from Avenella flexuosa and/or Nardus stricta. Only one specimen from the Czech Republic was known previously, collected in the Krkonoše Mts. (LAUTERER & NOVOTNÝ 1991). Based on recent collections, it is also reported here for the Krušné hory [Erzgebirge] Mts. in Bohemia (it also occurs on the German side of the Erzgebirge; e,g. NICKEL 2003, KUNZ et al. 2011) and, for the first time, for Moravia as well: the Jeseníky and Králický Snìžník Mts. (Fig. 13). The species is also known from the adjacent Polish side of the Sudetes Mts. (ŚWIERCZEWSKI & WALCZAK 2011b).

Cicadellidae: Cicadellinae

Cicadella lasiocarpae Ossiannilsson, 1981 (Figs 11, 25) Material examined. Bohemia: Horusice, 2 km SE, Ruda NNR (6854), 49°09′03″N, 14°41′27″E, 415 m a.s.l., 13.ix.2010, 20 ♀♀, I. Malenovský leg. (MMBC), 2 ♀♀, P. Kment leg. (NMPC).

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Fig. 13. Distribution of Anoscopus alpinus (Wagner, 1955) in the Czech Republic.

Remarks. Known from northern, western, and central Europe (Scandinavia, Russia, Great Britain, Ireland, Poland, Luxemburg and Germany), and the eastern Palaearctic: Siberia and Korea (NICKEL 2003, NIEDRINGHAUS et al. 2010, HOCH 2013). Feeding on Carex spp. and locally, although often abundantly, occurring in intermediate bogs, peat- bogs, peaty meadows and pastures, fens, and paludic lakeshores; Carex lasiocarpa, C. nigra and C. vesicaria have been explicitly mentioned as host plants (OSSIANNILSSON 1981, SZWEDO & GÊBICKI 1998, TISHECHKIN 2000, NICKEL 2003, SÖDERMAN 2007, KUNZ et al. 2011). The species is similar to the much more common, eurytopic and polyphagous (Linnaeus, 1758) from which it may be distinguished particularly by the colour pattern of the head and body, the size of the male apodemes and the serration of the female ovipositor (OSSIANNILSSON 1981, SZWEDO & GÊBICKI 1998, TISHECHKIN 2000, BIEDERMAN & NIEDRINGHAUS 2004). Despite the revision of a large number of Cicadella specimens from throughout the Czech Republic, only a series from a well-preserved intermediary bog in southern Bohemia, the Tøeboòsko Protected Landscape Area (Fig. 11), collected from Carex lasiocarpa and C. chordorrhiza, has proved to belong to C. lasiocarpae. New species for the Czech Republic.

246 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Cicadellidae: Typhlocybinae

Chlorita dumosa (Ribaut, 1933) (Fig. 11)

Published data. DLABOLA (1946): Podhoø near Praha (5852). DLABOLA (1954): Praha environs, Troja (5852). Material examined. Bohemia: Praha-Bohnice, Podhoøí (5852), 14.vii.1946, 6 ♂♂ 1 ♀ and v.1948, 1 ♂, all A. Hoffer leg., J. Dlabola det., I. Malenovský revid. (MNHN). Moravia: Znojmo-Bohumilice, 1 km SE, Naèeratický kopec Hill (7162), 48°50′12″N, 16°05′39″E, 250 m a.s.l., 26.vii.2009, dry and ruderal grassland in abandoned military training ground, 1 ♂ 2 ♀♀, I. Malenovský leg. (MMBC). Remarks. Living in cushions of Thymus spp. (Th. praecox, Th. serpyllum, and probably also Th. pulegioides) in sunny, dry to moderately dry, low-vegetated, usually grazed sites on sand, limestone and other well-drained substrates (NICKEL 2003). Quite widespread in Europe except the Iberian peninsula and the British Isles, reaching southern Scandinavia and the Baltic countries in the north and Kazakhstan in the east (NAST 1972, SÖDERMAN 2007, HOCH 2013). In Germany, the species is widespread although only local in the southern and central parts (NICKEL 2003). In Austria, it is known only from the eastern part (HOLZINGER 2009a). There is only one old record from the Czech Republic, from xerothermic slopes in the Vltava [Moldau] river valley in Prague (DLABOLA 1946) and the species was entered as “regionally extinct” in the Red List of Czech invertebrates (MALENOVSKÝ & LAUTERER 2005a). Its presence in the country is confirmed here by a first record from Moravia close to the Austrian border (Fig. 11); Ch. dumosa was recently collected in a patch of xerothermic grassland disturbed by vehicles in a former military training area.

Eupteryx lelievrei (Lethierry, 1874) (Fig. 14)

Published data. LAUTERER (1983): Bylnice, Bylnièky (6974); Knìždub, Èertoryje NNR (7170); Javorník, SW slopes of Háj hill (7171). MALENOVSKÝ & LAUTERER (2012): Valašské Klobouky, Javorùvky NR, Dobšená NM, Brumovka Brook valley, Na Nivách, and Bílé potoky NR (all 6874); Poteè, Ploštiny NR (6874); Nedašov, Kaòoury NM (6874); Bylnice, Lazy NR (6974); Hluk, Babí hora NM (7071); Vyškovec, Vyškovecké Bošaèky and Pod Hribovòou NM (7073); Knìždub, Knìždubský háj Forest and Èertoryje NNR (7170); Velká nad Velièkou, Zahrady pod Hájem NNR (7171); Javorník, Jazevèí NNR, Petruchovy Mlýny and Machová NR (7171); Nová Lhota, Vápenky, Porážky NNR (7171); Strání, Záhumenice NM (7172). Material examined. Bohemia: Maleèov, 3.5 km S, Babínské louky NM (5450), 50°35′57″N, 14°07′40″E, 530 m a.s.l., 28.viii.2013, species-rich meadows, 3 ♂♂. Lužnice, 1.2 km W, Velký a Malý Tisý NNR, intermittently wet Molinia meadow on edge of Šatlavy wetland (6954), 49°03′32″N, 14°44′21″E, 425 m a.s.l., 14.ix.2010, 1 ♀. All specimens I. Malenovský leg. (MMBC). Remarks. Monophagous on Betonica officinalis (Lamiaceae) in temporarily wet sites, e.g. along forest and shrub margins and in low-input meadows, preferentially on clayey soils (NICKEL 2003). Distributed mainly in central Europe, Belgium, the Pyrenees in France, Bulgaria and middle Russia, but generally rare and local (REMANE & FRÖHLICH 1994, NICKEL 2003). Recently found in southern Poland (ŒWIERCZEWSKI & WALCZAK 2011a). In the Czech Republic previously known only from species-rich, traditionally non-intensively managed meadows in the Bílé Karpaty Mts. [White Carpathians] in south-eastern Moravia, where the species is quite common (MALENOVSKÝ & LAUTERER

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Fig. 14. Distribution of Eupteryx lelievrei (Lethierry, 1874) in the Czech Republic.

2012). Here it is reported for the first time for Bohemia as well, in the Tøeboòsko Protected Landscape Area in southern Bohemia and the Èeské støedohoøí Protected Landscape Area in north-western Bohemia (Fig. 14).

Eupteryx signatipennis (Boheman, 1847) (Figs 15, 26)

Published data. MALENOVSKÝ (2006): Litice, Pod Hvìzdou NM (5352). Material examined. Bohemia: Želnava-Záhvozdí, 2 km W, wet meadows in Uhlíkovský potok Brook valley under Mt Èerný les (7149), 48°49′44″N, 13°58′43″E, 820 m a.s.l., 6.viii.2001, 1 ♂ 18 ♀♀. Èerná v Pošumaví, 2 km E, Slavkovické louky NM (7250), 48°44′01″N, 14°08′12″E, 760 m a.s.l., 21.viii.1998, wet Filipendula ulmaria grassland, 1 ♀. Horní Planá-Pøední Zvonková, 5.5 km S Horní Planá, Raèínská prameništì NM (7250), 48°43′16″N, 14°01′50″E, 750–770 m a.s.l., 27.viii.2001, 2 ♀♀. Moravia: Karlova Studánka, 5.5 km SW, Pradìd NNR, Velká kotlina Cirque (5969), 50°03′50″N, 17°14′10″E, 1200 m a.s.l., 17.viii.2011, subalpine tall- forb vegetation, on Filipendula ulmaria, 3 ♂♂ 21 ♀♀. All specimens I. Malenovský leg. (MMBC). Remarks. Monophagous on Filipendula ulmaria (Rosaceae) in wet, cool sites such as fens, along shores and ditches, and in disused meadows (NICKEL 2003). Quite widespread in northern, central and western Europe, absent from southern Europe with the exception of the Alps and the Pyrenees (REMANE & FRÖHLICH 1994, HOCH 2013). There is only one previous published record from the Czech Republic, from the Kokoøínsko Protected Landscape Area in northern Bohemia (MALENOVSKÝ 2006). The above-cited material

248 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Fig. 15. Distribution of Eupteryx signatipennis (Boheman, 1847) in the Czech Republic.

also documents its occurrence in the Šumava Mts. (Bohemian/Bavarian Forest) in southern Bohemia and, for the first time, from Moravia: the Jeseníky Mts. as well (Fig. 15). There are additional unpublished records from the Jizerské hory Mts. in northern Bohemia (P. Lauterer, pers. comm.). The species may be more widespread in the country but it is apparently absent from warm lowlands and has not yet been found in the Carpathians (cf. MALENOVSKÝ & LAUTERER 2012).

Zyginidia pullula (Boheman, 1845) (Fig. 16)

Published data. LAUTERER (1958): Brno-Holásky (6865). LAUTERER (1995): Lednice, Horní les (7166). MALENOVSKÝ & LAUTERER (2005b): Brno-Slatina (6866). MALENOVSKÝ et al. (2011): dry grassland sites in southern Moravia (many records in grids 6866, 6867, 6963, 6964, 6965, 7066, 7067, and 7266). MALENOVSKÝ & LAUTERER (2012): Bílé Karpaty Mts. (many records in grids 6874, 6974, 7071, 7072, 7073, 7170, 7171, and 7172). MALENOVSKÝ et al. (2013): Mikulov, Svatý kopeèek NR (7165); Sedlec, Slanisko u Nesytu NNR (7266). Material examined. Bohemia: Žehuò, 2 km NE, Žehuòský rybník NNM, intermittently to permanently wet meadows on N shore of Žehuòský rybník Pond (5857), 50°09′08″N, 15°18′20″E, 198 m a.s.l., 22.viii.2012, 1 ♂, I. Malenovský leg. Moravia: Prosetín (6464), field of Trifolium pratense and Lolium multiflorum, 600 m a.s.l., 15.x.1999, 1 ♀, I. Malenovský leg. Tišnov, Kvìtnice NR, SE slopes (6664), 360–400 m a.s.l., 17.ix.1997, dry grassland on limestone, 1 ♂ 1 ♀, P. Lauterer leg. et det. Rájeèko, 1 km W (6665), field of Trifolium pratense and Lolium multiflorum, 400 m a.s.l., 10.vi.1998, 1 ♀, 6.vii.1998, 1 ♂, 29.vii.1998, 3 ♂♂ 2 ♀♀, 2.ix.1998, 3 ♂♂, 22.ix.1998, 2 ♂♂ 2 ♀♀, 10.x.1998, 2 ♂♂ 4 ♀♀, all I. Malenovský leg. Blansko-Klepaèov (6665), field

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Fig. 16. Distribution of Zyginidia pullula (Boheman, 1845) in the Czech Republic.

of Trifolium pratense and Lolium multiflorum, 340 m a.s.l., 25.ix.1998, 2 ♂♂ 1 ♀, 10.x.1998, 1 ♂ 3 ♀♀, 25.vi.1999, 9 ♂♂ 3 ♀♀, 25.vii.1999, 4 ♂♂ 3 ♀♀ 1 nymph, 25.ix.1999, 21 ♂♂ 21 ♀♀, 12.x.1999, 30 ♂♂ 17 ♀♀, all I. Malenovský leg. Vranov u Brna towards Brno-Útìchov (6665), 450 m a.s.l., 3.xi.1996, forest undergrowth, 4 ♀♀, P. Lauterer leg. et det. Domašov (6764), 460 m a.s.l., red clover field, 21.ix.1998, 1 ♂ 1 ♀, and 21.x.1998, 3 ♀♀, I. Malenovský leg. Brno-Knínièky, shores of the Brnìnská pøehrada Dam near Osada (6765), 230 m a.s.l., 9.viii.2009, 1 ♂, I. Malenovský leg. Brno-Komín, Sítí (6765), 8.viii.2009, 250 m, abandoned orchard, 1 ♂, I. Malenovský leg. Brno-Obøany, Malomìøický lom Quarry (6765), 220 m a.s.l., viii.2008, 1 ♂, P. Baòaø leg. Brno-Malomìøice, Hády Hill, Mìstský lom and Rùženin lom Quarries (6766), 370–400 m a.s.l., 8.ix.2008, 4 ♂♂ 3 ♀♀, and 22.ix.2009, several ♂♂ ♀♀, I. Malenovský leg. Senorady, Údolí Oslavy a Chvojnice NR, slopes under Levnov (= Ketkovický hrad) castle ruins (6863), 300–400 m a.s.l., 11.v.2008, 3 ♂♂ 1 ♀, I. Malenovský leg. Mohelno, Mohelenská hadcová step NNR (6863), 310–340 m, 31.v.1996, many ♂♂ ♀♀, P. Lauterer leg. et det. Brno-Chrlice, along Dvorský potok Brook (6865), 205 m a.s.l., 20.viii.2008, 1 ♀, P. Baòaø leg. Brno-Chrlice, Mokøina u dálnice wetland (6865), 200 m a.s.l., 9.ix.2008, 1 ♂, I. Malenovský leg. Brno-Chrlice, environs of Splavisko Ponds (6865), 9.ix.2008, 1 ♂, I. Malenovský leg. Brno- Holásky, Holásecká pískovna Sandpit (6865), 220 m a.s.l., 9.ix.2008, 1 ♀, I. Malenovský leg. Brno-Starý Lískovec, abandoned field at Leskava Brook (6865), 230 m a.s.l., viii.2008, 1 ♀, P. Baòaø leg. Brno-Líšeò, Pod Oøíšky, orchards (6866), viii.2008, 280 m a.s.l., 1 ♂, P. Baòaø leg. Jamolice, 2 km N (6963), 49°05′09″N, 16°15′29″E, 370 m a.s.l., 2.viii.2010, grassland and open woods in abandoned military training ground, 1 ♂, I. Malenovský leg. Výrovice, 0.5 km S, Výrovické kopce Hills (7062), 270 m a.s.l., 7.viii.2009, 1 ♂, P. Baòaø & I. Malenovský leg. Bzenec, 1 km SE, sands (7069), 190-195 m, 17.viii.1995, 1 ♂ 1 ♀, P. Lauterer leg. Znojmo- Bohumilice, 1 km SE, Naèeratický kopec Hill (7162), 48°50′12″N, 16°05′39″E, 250 m a.s.l., 26.vii.2009, dry and ruderal grassland in abandoned military training ground, 1 ♀, I. Malenovský leg. All material in MMBC. Remarks. Feeding on various grasses (Poaceae), e.g. Bromus, Cynodon, Digitaria,

250 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Echinochloa, Elymus, Molinia, Setaria, and cultivated cereals and maize, to which occasional damage has been reported from Italy (VIDANO & ARZONE 1985). A complete life-cycle was observed in the Czech Republic on Lolium multiflorum, but the species definitely has at least some additional grass species as local hosts. Widely distributed from Mongolia, Kazakhstan and Turkey to south-eastern and central Europe, southern Sweden and Denmark, and north-eastern Spain (REMANE & FRÖHLICH 1994), but apparently absent from some areas, e.g. from faunistically well-explored Germany, there is only one record of a single specimen from Bavaria (REMANE & FRÖHLICH 1994, NICKEL 2003). Recently, Z. pullula has been recorded from southern Poland (ŒWIERCZEWSKI & WALCZAK 2011a). In the Czech Republic, Z. pullula is widespread and common in southern Moravia, where it lives in various kinds of dry to moderately wet grassland: meadows, pastures, xerothermic grassland, forest glades, ruderal sites, and fields from the lowlands up to at least 700 m (MALENOVSKÝ & LAUTERER 2005b, 2012; MALENOVSKÝ et al. 2011), while it appears to be much rarer or absent in large parts of Bohemia: the first record for the latter part of the country is presented here (Fig. 16). The distribution of Z. pullula in the River Elbe lowlands in central Bohemia overlaps with that of the closely-related Z. scutellaris (Herrich-Schäffer, 1838), a species with much the same ecology, expanding its range in the Czech Republic in recent years and, in contrast to Z. pullula, not yet recorded from Moravia (MALENOVSKÝ & TROPEK 2009, TROPEK et al. 2012, and additional recent unpublished data in MMBC). Zyginidia pullula has two generations per year in the Czech Republic and overwinters in the egg stage, which has been confirmed by dissecting the ovaries of females from samples throughout the season. Adults of the first generation occur from the second decade of May to the end of July, with the second generation from the beginning of August to the end of October; ripe eggs have been found in the ovaries of females of the first generation from the last decade of June to July and for the second generation in the second half of October. In Italy, three generations with adults overwintering are known (VIDANO & ARZONE 1985) while in Finland there is only one generation per year and the species overwinters in the egg stage (SÖDERMAN 2007).

Cicadellidae: Deltocephalinae

Allygidius abbreviatus (Lethierry, 1878) (Fig. 17)

Published data. DLABOLA (1956): Pavlovské vrchy Hills (7165). MALENOVSKÝ et al. (2011): Újezd u Brna, Špice NR (6866); Køižanovice near Buèovice, Èlupy NR (6867); Rašovice, Rašovický zlom-Chobot NR (6867); Dolní Dubòany, Ve Žlebì (6963); Syrovice, Bezourek (6965); Bluèina, Nové hory NR (6965); Kurdìjov, Kamenný vrch u Kurdìjova NR (7066); Hustopeèe, Pøední kopaniny (7066); Nìmèièky, Straèí (7066); Úvaly near Valtice, Kameníky (7266). MALENOVSKÝ & LAUTERER (2012): Poteè, Poštiny NR (6874); Valašské Klobouky, Bílé potoky NR (6874); Nedašov, Jalovcová stráò NR (6874); Brumov, Kloboucká street (6974); Blatnièka, Milejovské louky (7071); Horní Nìmèí, Drahy NR (7071); Radìjov, Žerotín NM (7169); Knìždub, Èertoryje NNR (7170); Radìjov, Veselka hill (7170); Velká nad Velièkou, Zahrady pod Hájem NNR (7171); Suchov, Trnovský Mlýn (7171); Javorník, Jazevèí NNR, Petruchovy Mlýny and Machová NR (7171); Nová Lhota, Fojtické Mlýny, Vápenky, and Porážky NNR (7171). Material examined. Bohemia: Blšany u Loun, 0.7 km W, Blšanský chlum Hill (5649), 50°20′44″N, 13°50′20″E, 260 m a.s.l., 1.viii.2009, thermophilous fringe and open pine wood, 1 ♀, I. Malenovský leg.;

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Fig. 17. Distribution of Allygidius abbreviatus (Lethierry, 1878) in the Czech Republic.

29.viii.2009, 1 ♀, O. Èížek & P. Marhoul leg. Moravia: Rozhraní, southern part of village, towards Dolní Smržov (6465), 380–430 m, 7.vii.1987, 1 ♂, P. Lauterer leg. Malhostovice, Malhostovická pecka NM (6664), 350 m, 19.vii.1998, 1 ♂ 1 ♀, P. Lauterer leg. et det. Køtiny, arboretum (6666), 440–450 m a.s.l., 16.viii.1978, 1 ♀, L. Pospíšilová leg. Pøibyslavice-Devìt køížù (6763), 500 m a.s.l., 10.vii.1997, forest undergrowth, 1 ♂, P. Lauterer leg. et det. Brno-Komín, Komínský vršek Hill (6765), 28.vi.2009, 275 m, 1 ♂, I. Malenovský leg. Brno-Komín, Sítí (6765), 28.vi.2009, 250 m, abandoned orchard, 1 ♂, I. Malenovský leg. Brno-Komín, Netopýrky NM (6765), 280 m, 30.vi.2012, 1 ♀, I. Malenovský leg. Brno-Medlánky, Medlánecká skalka NM (6765), 280 m, 30.vi.2012, 1 ♀, I. Malenovský leg. Brno-Malomìøice, Hádecká planinka NNR (6766), 400–420 m, 9.vii.1997, 4 ♂♂ 5 ♀♀, P. Lauterer leg. et det. Mohelno, Mohelenská hadcová step NNR (6863), 310–340 m, 10.vii.1997, 1 ♂, P. Lauterer leg. et det. Rebešovice, 1 km NE, Velké Družïavy NM (6865), 220 m a.s.l., 24.vii.1978, 1 ♂ 1 ♀, J. L. Stehlík leg. Ostopovice (6865), railway embankment, 230 m, 7.ix.2004, 1 ♀, I. Malenovský leg. Žarošice (6967). ca. 250 m a.s.l., 8.viii.1962, 1 ♀, J. L. Stehlík leg. Archlebov, Dubový vrch hill (6968), 250–300 m, 24.vi.2000, 1 ♀, P. Lauterer leg. Výrovice, 0.5 km S, Výrovické kopce Hills (7062), 270 m a.s.l., 7.viii.2009, 1 ♀, P. Baòaø & I. Malenovský leg. Podmolí-Šobes (7161), 320 m, 12.vii.1997, vineyard and oak wood, 1 ♀, P. Lauterer leg. et det. Znojmo-Naèeratice, 1.5 km NW, Naèeratický kopec Hill (7162), 270 m a.s.l., 26.vii.2009, grassland and open woods in abandoned military training ground, 4 ♀♀, I. Malenovský leg. Klentnice (7165), 0.3 km W of village, 400 m, 31.viii.2004, 1 ♀, I. Malenovský leg. Klentnice, Tabulová NNR (7165), 400–450 m a.s.l., 9.vii.1968, 3 ♂♂ 1 ♀, P. Lauterer leg. et det. Mikulov, 0.5 km E (7165), 250–260 m, 26.vi.2001, vineyard, 1 ♂, P. Lauterer leg. et det. Sedlec, Kienberg NM (7166), 210–230 m, 19.vi.2010, 2 ♂♂, I. Malenovský leg. Sedlec, Lišèí vrch NR (7266), 280–300 m, 19.vi.2010, 1 ♂, I. Malenovský leg. All material in MMBC. Remarks. In the Czech Republic, usually collected among tall grass in non-intensively used meadows and pastures on relatively dry, warm and sunny sites, in fragments of

252 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic xerothermic grassland, and along forest margins (MALENOVSKÝ & LAUTERER 2012). Nymphs develop in the herbal layer while adults may also be found on trees, e.g. cherry (HEGAB et al. 1980). Definite host plant records are lacking; the species has been collected on places with dominant Arrhenatherum elatius, Bromus erectus, Brachypodium pinnatum, Calamagrostis epigejos, and Molinia arundinacea, at least some of which may prove to be larval hosts. It is also distributed in southern parts of central Europe, northern France, Belgium, Italy, Bulgaria, and Tunisia (NICKEL 2003, HOCH 2013). The record from Tunisia was published by MELICHAR (1899); the corresponding voucher specimens could be located in the Melichar collection in MMBC (2 ♀♀ with the label “Tunis”) but their true origin may be doubtful, as in the case for several other Auchenorrhyncha species allegedly collected in Tunisia by German 19th- century entomologist O. Schmiedenknecht (R. Remane and H. Nickel, pers.comm.). In the Czech Republic, A. abbreviatus is quite common in southern Moravia but was not previously known from Bohemia – first recorded here for this part of the country from a dry and warm region in north-western Bohemia (Fig. 17).

Euscelis ohausi Wagner, 1939 (Figs 11, 27) Material examined. Bohemia: Podboøany-Valov, 0.5 km W (5746), 50°12′08″N, 13°24′29″E, 350 m a.s.l., 2.viii.2009, heathland in abandoned military training ground, on Cytisus scoparius, 2 ♂♂ 3 ♀♀, I. Malenovský leg. (MMBC). Remarks. A western European species known from Portugal, France, Switzerland, Germany, Belgium, the Netherlands, Britain, Denmark and north-western Poland (NICKEL 2003, HOCH 2013) and recently introduced into North America (HOEBEKE & WHEELER 2010). It lives mostly in sunny, damp to moderately dry sites on acidic substrates (NICKEL 2003). There are two forms in central Europe which differ in host plant, colouration, body size and distribution: f. typica lives on Genista anglica e.g. near the coast of the North Sea and the Baltic Sea in Germany while f. singeri Wagner, 1951 dwells on Cytisus scoparius and is known e.g. from central and southern Germany and Luxemburg (NICKEL 2003, NIEDRINGHAUS et al. 2010, KUNZ et al. 2011). The Czech specimens found in north-western Bohemia (Fig. 11), to judge by morphology and host plant, belong to f. singeri. Both forms are treated as synonyms at species level by some authors (e.g. OSSIANNILSSON 1983) as they can interbreed and produce fertile offspring (STRÜBING 1978) or as distinct subspecies (e.g. NICKEL 2003). New species for the Czech Republic.

Fieberiella septentrionalis Wagner, 1963 (Fig. 18)

Published data. DLABOLA (1965): Moravia (historical specimens without more definite locality data). MALENOVSKÝ et al. (2011): Ivanèice-Budkovice, Krumlovsko-Rokytenské slepence NNR (Budkovické skály) (6964). Material examined. Bohemia: Kadaò-Prunéøov, 1 km E, “Starý lom” ( ash deposits of Prunéøov power- plant; 5545), 50°24′59.687″N, 13°16′29.125″E, ca. 350 m a.s.l., yellow pan traps, 30.–31.vii.2010, 1 ♀, 20.–22.viii.2010, 1 ♂, R. Tropek leg., I. Malenovský det. Plzeò-city (6246), grounds of school of agriculture,

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Fig. 18. Distribution of Fieberiella septentrionalis Wagner, 1963 in the Czech Republic.

9.viii.1999, Galinsoga parviflora, 1 ♂, L. Sitaøová leg., P. Lauterer det. Èeský Krumlov-Nové Dobrkovice and Vyšný, Vyšenské kopce NNR (7151), 48°49′18″N, 14°17′41″E, 530 m a.s.l., 2.viii.1998, 2 ♂♂, I. Malenovský leg.et det. (all material MMBC). Remarks. Polyphagous on various shrubs, trees and tall herbs, frequently associated with Prunus spinosa, Rosa spp., Ligustrum spp. and other species in dry grassland, ruderal sites, along forest margins and hedges but also synanthropic in gardens and parks in Germany (MEYER-ARNDT & REMANE 1992, NICKEL 2003). Widely distributed from central Europe through the eastern part of the Balkan peninsula, North Anatolia and central Asia to north-east China (MEYER-ARNDT & REMANE 1992). In the Czech Republic probably widely distributed but only poorly documented: known to date only from Moravia (DLABOLA 1965, MALENOVSKÝ et al. 2011). Reported here for the first time for Bohemia (Fig. 18).

Jassargus allobrogicus (Ribaut, 1936) (Fig. 19)

Published data. DLABOLA (1944, 1954): Peøimov near Jilemnice (5358), Kozákov Hill near Semily (5457), Praha-Modøany (5952), Praha-Radotín (6052). KULA (2002): Jílové-Snìžník – Kristin Hrádek, Letadlo, Ostrov, Vlèák (5150), Snìžník (5250); Tisá (5250). Material examined. Bohemia: Pøebuz, 1 km NE, Pøebuzské vøesovištì NM (5641), 880 m a.s.l., 28.ix.2012, secondary montane heath on former peat diggings, 1 ♂ 3 ♀♀, I. Malenovský leg. Pøebuz, 2.8 km NW, Rolavská

254 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Fig. 19. Distribution of Jassargus allobrogicus (Ribaut, 1936) in the Czech Republic.

vrchovištì NNR, Lieche (= Rolavský rybník Pond) (5641), 910 m a.s.l., 21.vi.2011, spruce forest edge, 1 ♂ 1 ♀, I. Malenovský leg.; 12.viii.2011, 1 ♂ 2 ♀♀, P. Kment leg.; 29.ix.2012, 1 ♂, I. Malenovský leg. Pøebuz, 3.6 km NW, Rolavská vrchovištì NNR, Velký cínový dùl (= former mine Rolava or Sauersack) (5641), 930 m a.s.l., 21.vi.2011, heath, 1 ♂, I. Malenovský leg.; 11.viii.2011, 2 ♂♂ 4 ♀♀, P. Kment leg.; 28.ix.2012, 2 ♂♂ 2 ♀♀, I. Malenovský leg. Pøebuz, 3 km W, former village of Chaloupky (5641), 20.–22.vi.2011, 830 m, submontane grassland, 2 ♂♂, I. Malenovský leg. Prameny, 2 km NE, Køížky NNM (5942), 50°03′57″N, 12°44′59″E, 800 m a.s.l., 31.vii.2007, dry to mesic grassland on serpentinite rocks, 2 ♂♂ 1 ♀, I. Malenovský leg. Moravia: Kobylá nad Vidnavkou, 2 km E, NE slope of Smolný vrch Hill near Venušiny misky NNM (5668), 50°20′11″N, 17°09′07″E, 350 m a.s.l., 18.viii.2011, forest clearing, on Avenella flexuosa, 12 ♂♂ 3 ♀♀, I. Malenovský leg. Karlova Studánka, 6 km SW, Pradìd NNR, N slope of Mt Petrovy kameny (5969), 50°04′20″N, 17°13′52″E, 1350–1400 m a.s.l., 17.viii.2011, open montane spruce forest and subalpine grassland, 1 ♂, I. Malenovský leg. (MMBC). Remarks. In open deciduous and coniferous forests, and dry pastures and heaths, especially common on acidic substrates on Avenella flexuosa under beech, spruce and pine but probably also feeding on fine-leaved Festuca spp. (NICKEL 2003, SÖDERMAN 2007). Widespread in western, central and northern Europe, eastward to central Russia (HOCH 2013). In the Czech Republic previously known only from northern and central Bohemia: several records are added here from the Krušné hory [= Erzgebirge] and the Slavkovský les Mts. in western Bohemia and the Hrubý Jeseník Mts. and the Vidnavsko region in Silesia (northern Moravia). The species thus appears to be widespread at least in the northern half of the country, from hills to subalpine elevations, but has not been

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Fig. 20. Distribution of Recilia coronifer (Marshall, 1866) in the Czech Republic.

found in large areas with relatively warm climate and basic substrates, such as southern Moravia (Fig. 19).

Pinumius areatus (Stål, 1858) (Figs 11, 28)

Published data. LAUTERER (1983): Vacenovice, S margin of village (7069). Material examined. Bohemia: Kadaò-Tušimice, “Téèko” (fly ash deposits of Tušimice power-plant; 5646), 50°22′25″N, 13°20′59″E, ca. 310 m a.s.l., yellow pan traps, 5.–6.vi.2010, 22 ♂♂ 7 nymphs, 30.–31.vii.2010, 5 nymphs, 20.–22. viii.2010, 35 ♂♂ 6 ♀♀ 3 nymphs, all R. Tropek leg., voucher specimens in MMBC. Remarks. Widely distributed in central and eastern Europe, Kazakhstan, Kirghizia, Siberia, Mongolia and North America (NAST 1972, OSSIANNILSSON 1983, HOCH 2013). Reported to live in sunny, sparsely vegetated dry grassland, usually on aeolian or moraine sands on grasses (OSSIANNILSSON 1983, NICKEL 2003). The host plant in central Europe is Festuca ovina (KUNZ et al. 2011). In central Europe, P. areatus is an extremely rare and local species. The last records from Austria date back to 1940 and the species has not been found since (HOLZINGER 2009b). In Germany, there are a few recent sites in coal- mining and sand-quarrying areas in eastern Germany (NICKEL 2003, H. Nickel pers. comm.). From the Czech Republic, there is only one finding of two specimens in an area of loose sands in south-eastern Moravia, dating back to 1963 (LAUTERER 1983). As this

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Moravian locality, near Vacenovice, was later destroyed, P. areatus has been considered an extinct species in the Czech Republic (LAUTERER 1983, MALENOVSKÝ & LAUTERER 2005a). Its presence in the country is confirmed here based on recent first records from Bohemia: a strong population of P. areatus was discovered during a sampling of the fauna of the fly-ash deposits of a large coal-fired power station in north-western Bohemia (Fig. 11). This extreme, post-industrial habitat has recently been shown to provide refuge for several critically endangered insects, particularly those originally confined to inland sand dunes (TROPEK et al. 2013). The morphology of the Czech specimens, particularly the shapes of the male pygophore appendage and aedeagus, closely matches illustrations of P. areatus given by EMELJANOV (1972) and OSSIANNILSSON (1983) and differs from the diagnosis of the central-Asian/Siberian species Pinumius occultus Emeljanov, 1966 and P. nebulicola Emeljanov, 1972 (see HAMILTON 1983, REMANE 1987 and NICKEL 2003 for a discussion on the taxonomic identity of European Pinumius populations).

Recilia coronifer (Marshall, 1866) (Figs 20, 29)

Material examined. Bohemia: Sobotka (5557), 20.vii.1941, forest clearing, 1 ♂, J. Dlabola leg. et det., I. Malenovský revid. (MNHN). Dlouhopolsko/Knìžièky, meadows and woods in the environs of the Dlouhopolský rybník and Kopièácky rybník Ponds (5857), 50°09′40″N, 15°19′60″E, 240 m a.s.l., 22.viii.2012, 1 ♀, I. Malenovský leg. (MMBC). Moravia: Jamolice, 2 km N (6963), 49°05′09″N, 16°15′29″E, 370 m a.s.l., 2.viii.2010, grassland and open woods in abandoned military training ground, 2 ♂♂ 2 ♀♀, I. Malenovský leg. (MMBC). Znojmo-Naèeratice, 1.5 km NW, Naèeratický kopec Hill (7162), 48°49′40″N, 16°05′42″E, 270 m a.s.l., 26.vii.2009, grassland and open woods in abandoned military training ground, 1 ♂, I. Malenovský leg. (MMBC). Remarks. Mainly in open oak and pine forests, clearings, along forest margins and roads, bog margins and in meadows on Holcus mollis and Molinia caerulea agg. (NICKEL 2003). Widespread from the Iberian peninsula and Great Britain in the west through central Europe, the Balkan peninsula and the Near East to the eastern Palaearctic (HOCH 2013) but so far insufficiently documented in the Czech Republic. DLABOLA (1943) probably recorded the above-cited specimen from Sobotka as “Thamnotettix coroniceps Kbm. 1868” but later listed it in his monograph on the Czechoslovak fauna under “ schmidtgeni (Wagner, 1939)” together with data from Moravia (Èejè and Hodonín) and Slovakia (Chotín and Štúrovo), apparently treating the two species names as synonyms (DLABOLA 1954). In the check-list of the Auchenorrhyncha of Czechoslovakia, DLABOLA (1977) included only two Recilia species (R. horvathi (Then, 1896) from Moravia and Slovakia, and R. schmidtgeni from Bohemia, Moravia and Slovakia). Corresponding specimens of Recilia Edwards, 1922 and Distant, 1917 (see WEBB & VIRAKTAMATH 2009 for currently valid nomenclature) in the Dlabola collection in MNHN have been revised recently: as well as a single specimen of R. coronifer from Bohemia (Sobotka), there were two males of Maiestas schmidtgeni (Wagner, 1939) from Moravia (Èejè, viii.1940, 1 ♂ and 30.viii.1942, 1 ♂, both A. Hoffer leg.) and two males of M. schmidtgeni from Slovakia (Bratislava, 1945, 1 ♂, Görtler leg.; Parkán (= Štúrovo), vii.1947, 1 ♂, A. Hoffer leg.). Maiestas horvathi is represented in the

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Dlabola collection only by specimens from Moravia (Brno-Knínièky, Brnìnská pøehrada Reservoir, 18.vi.1954, 1 ♂; Brno, at light, 1.ix.1955, 1 ♂; both P. Lauterer leg.). Based on this and the additional, recently-collected material cited above, R. coronifer is confirmed/recorded here for the Czech Republic for the first time. As far as is known, the species is present but uncommon in central/north-eastern Bohemia and south-western Moravia (Fig. 20). Currently, there is no confirmed record of M. schmidtgeni from Bohemia, in contrast to confirmed historical data from south-eastern Moravia. Maiestas horvathi is also known to occur in the Czech Republic only in southern Moravia (MALENOVSKÝ & LAUTERER 2005b, MALENOVSKÝ et al. 2011).

Acknowledgements I dedicate this paper to my teacher, Pavel Lauterer, who introduced me to entomology and, for many years, has generously shared with me his amazingly deep and universal knowledge and enthusiasm for the study of leafhoppers, planthoppers and psyllids, for which I am most grateful. I also thank Michal Horsák, Jan Roleèek and Jan Sychra (all from the Institute of Botany and Zoology, Masaryk University, Brno) for the organization of wonderful collecting excursions to various interesting regions of the Czech Republic, through which several taxa addressed in this paper could be recorded. Petr Kment is to be thanked for valuable assistance in the field on certain excursion trips and gifts of material, as well as for the hospitality and access to the collection of the National Museum, Prague. Pavel Marhoul (Daphne ÈR, Praha) and Oldøich Èížek (Biology Centre of the Czech Academy of Sciences and University of South Bohemia, Èeské Budìjovice) are acknowledged for their help in organization of the collecting trips to disused military training grounds that were supported by project VaV MŽP SP2/d3/153/08. Robert Tropek (Biology Centre of the Czech Academy of Sciences and University of South Bohemia, Èeské Budìjovice) kindly provided Auchenorrhyncha material from fly-ash deposits from power stations in north-western Bohemia collected during a project supported by the Czech Science Foundation (P504/12/2525). A revision of specimens in the Jiøí Dlabola collection in the Muséum national d’Histoire naturelle in Paris was possible thanks to the hospitality of Adéline Soulier-Perkins and Thierry Bourgoin and a grant for invited scientists provided by MNHN in 2011. Dmitry Tishechkin (Moscow, Russia) is thanked for the information on the Czech specimens of Hephathus achilleae that he examined. Last but not least, I am obliged to Sabine Walter (Kurort Hartha, Germany) for a loan of specimens of Javesella simillima from her collection, Herbert Nickel (Göttingen, Germany) for critical reading of the manuscript and useful comments, Lenka Hubáèková and Jan Sychra (both Brno) for photographs of collecting sites and Tony Long (Svinošice) for helping to work up the English. This paper appears through financial support provided to the Moravian Museum, Brno by the Ministry of Culture of the Czech Republic as part of its long-term conceptual development programme for research institutions (ref. MK000094862).

258 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

Figs 21–29. Habitus of some Auchenorrhyncha species recorded from the Czech Republic. 21 – Kelisia confusa Linnavuori, 1957; 22 – Litemixia pulchripennis Asche, 1980; 23 – Hephathus achilleae Mityaev, 1967, in dorsal and frontal views; 24 – Anoscopus alpinus (Wagner, 1955); 25 – Cicadella lasiocarpae Ossiannilsson, 1981; 26 – Eupteryx signatipennis (Boheman, 1847); 27 – Euscelis ohausi Wagner, 1939; 28 – Pinumius areatus (Stål, 1858); 29 – Recilia coronifer (Marshall, 1866). Scale bars: 1 mm.

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References

ASCHE M. 1980: Litemixia pulchripennis gen. et spec. nov., eine neue Delphacide aus Südwest-Frankreich (Homoptera Cicadina Delphacidae). Marburger Entomologische Publikationen 1(3): 59–92. BEZDÌK J. 2011: Pøehled živoèišných druhù popsaných z území Èeské republiky. [A review of animal species described from the territory of the Czech Republic.] Mendelova univerzita v Brnì, Brno, 420 pp + CD ROM (in Czech, English summary). BIEDERMANN R. & NIEDRINGHAUS R. 2004: Die Zikaden Deutschlands. Bestimmungstafeln für alle Arten. Wissenschaftlich Akademischer Buchvertrieb-Fründ, Scheeßel, 409 pp. BOGUSCH P., STRAKA J. & KMENT P. (eds.) 2007: Annotated check-list of the Aculeata (Hymenoptera) of the Czech Republic and Slovakia. Komentovaný seznam žahadlových blanokøídlých (Hymenoptera: Aculeata) Èeské republiky a Slovenska. Acta Entomologica Musei Nationalis Pragae, Supplementum 11: 1–300. DANÈÁK M., DUCHOSLAV M. & TRÁVNÍÈEK B. 2012: Taxonomy and cytogeography of the Molinia caerulea complex in central Europe. Preslia 84: 351–374. DLABOLA J. 1943: Nìkolik zajímavých køískù z Èech a Moravy. (Einige interessante Zikadiden aus Böhmen und Mähren.) Entomologické listy (Brno) 6: 1–2 (in Czech, German summary). DLABOLA J. 1944: IV. pøíspìvek k poznání fauny køísù (Homopt. Auchenorrh.). IV. attributio ad cognitionem homopterorum faunae. Èasopis Èeskoslovenské spoleènosti entomologické 41: 94–100 (in Czech, Latin summary). DLABOLA J. 1946: Popisy 2 nových druhù køísù z Èech a jiné význaèné nálezy z území ÈSR. (Homopt. Auchenorrhyncha). Description de deux nouvelles espèces et plusieurs remarques sur les espèces peu connues d’Europe centrale. (Homopt. Auchenorrh). Acta Entomologica Musei Nationalis Pragae 24: 97–106 (in Czech, French summary). DLABOLA J. 1954: Køísi – Homoptera. Fauna ÈSR, vol. 1. [Leafhoppers and planthoppers – Homoptera. Fauna of Czechoslovakia, vol. 1.] Nakladatelství Èeskoslovenské akademie vìd, Praha, 338 pp (in Czech, German and Russian summary). DLABOLA J. 1956: Faunistika a nìkteré nové druhy palearktických køísù. Faunistik und neue Arten der palearktischen Zikaden (Homoptera, Auchenorrhyncha). Acta Entomologica Musei Nationalis Pragae 30(1955): 121–128 (in German, Czech introduction). DLABOLA J. 1965: Zoogeographische Arten-Gliederung der Gattung Fieberiella Sign. (Homopt., Auchenorrhyncha). Acta Entomologica Bohemoslovaca 62: 428–442. DLABOLA J. 1977: Homoptera Auchenorrhyncha. Pp. 83–96. In: DLABOLA J. (ed.): Enumeratio insectorum bohemoslovakiae. Checklist tschechoslowakische Insektenfauna. Acta Faunistica Entomologica Musei Nationalis Pragae, Supplementum 4: 1–159. DUDA L. 1892: Hmyz polokøídlý (Rhynchota) (Heteroptera, Cicadina, Psyllidae) v Èechách žijící. [Hemiptera (Rhynchota) (Heteroptera, Cicadina, Psyllidae) living in Bohemia.] Catalogus insectorum faunae bohemicae. I. Spoleènost pro fysiokracii, Praha, 44 pp (in Czech). EHERENDORFER F. & HAMANN U. 1965: Vorschläge zu einer floristischen Kartierung von Mitteleuropa. Berichte der Deutschen Botanischen Gesselschaft 78: 35–50. EMELJANOV A. F. 1972: Novye palearkticheskie tsikadki podsemeystva Deltocephalinae (Homoptera, Cicadellidae). (New Palaearctic leafhoppers of the subfamily Deltocephalinae (Homoptera, Cicadellidae).) Entomologicheskoe Obozrenie 51: 102–111 (in Russian, English title). EMMRICH R. 2003: History of Auchenorrhyncha research in central Europe. Geschichte der Zikadenkunde in Mitteleuropa. Pp. 5–25. In: HOLZINGER W. E., KAMMERLANDER I. & NICKEL H.: The Auchenorrhyncha of Central Europe. Die Zikaden Mitteleuropas. Volume 1: Fulgoromorpha, Cicadomorpha excl. Cicadellidae. Brill, Leiden – Boston, 673 pp. GIUSTINA W., DELLA & REMANE R. 1992: La Faune de France des Delphacidae. II. Notes de chasses faites, pour l’essentiel, en 1990 (Homoptera Auchenorrhyncha). Bulletin de la Société entomologique de France 96(1991): 313–330. HAMILTON K. G. A. 1983: Introduced and native leafhoppers common to the Old and New World (Rhynchota: Homoptera: Cicadellidae). Canadian Entomologist 115: 473–511. HEGAB A.M., OROSZ A. & JENSER G. 1980: Observations on the larvae and imagoes of some Allygus species (Homoptera). Folia entomologica hungarica 41: 61–66. HOCH H. 2013: Fauna Europaea: Hemiptera: Cicadomorpha etc. In: Fauna Europaea, version 2.6.1. Available online at http://www.faunaeur.org (accessed 4 July 2013).

260 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

HOEBEKE E. R. & WHEELER A. G. JR. 2010: Euscelis ohausi Wagner (Hemiptera: Cicadellidae: Deltocephalinae): a Palearctic leafhopper established in North America. Proceedings of the Entomological Society of Washington 112: 517–525. HOLZINGER W. E. 2009a: Checklisten der Fauna Österreichs, No. 4: Auchenorrhyncha (Insecta). Biosystematics and Ecology Series 26: 41–100. HOLZINGER W. E. 2009b: Rote Listen gefährdeter Zikaden (Auchenorrhyncha) Österreichs. Pp. 41–317. In: WALLNER R. & ZULKA K. P. (eds.): Rote Listen gefährdeter Tiere Österreichs Vol. 14(3). Böhlau, Wien, 534 pp. HOLZINGER W. E., KAMMERLANDER I. & NICKEL H. 2003: The Auchenorrhyncha of Central Europe. Die Zikaden Mitteleuropas. Volume 1: Fulgoromorpha, Cicadomorpha excl. Cicadellidae. Brill, Leiden – Boston, 673 pp. HULDÉN L. 1974: The Javesella discolor group (Homoptera, Delphacidae) of North Europe, with description of a new species. Notulae Entomologicae 54: 114–116. JEŽEK J. 1994: RNDr. Jiøí Dlabola, CSc., sedmdesátiletý. 70th Anniversary celebrated by RNDr. Jiøí Dlabola, CSc. Èasopis Národního Muzea, Øada pøírodovìdná 163: 69–77 (in Czech, English title). KMENT P. 2009: Èechy a Morava pro potøeby faunistického výzkumu. Bohemia and Moravia for the purposes of faunistic research. Klapalekiana 45: 287–291 (in Czech, English summary). KOLENATI F. A. 1859: Naturhistorische Durchforschung des Altvatergebirges. Jahresheft der naturwissen- schaftlichen Section der k.k. mähr. schles. Gesellschaft für Ackerbau, Natur- und Landeskunde, für das Jahr 1858 1859(1858): 1–83. KOLENATI F. A. 1860: Einige neue Insekten-Arten vom Altvater (dem hohen Gesenke der Sudeten). Wiener Entomologische Monatschrift 4: 381–394. KOLEŠKA Z. 1998: Seznam biografií èeskoslovenských entomologù (entomologové nežijící). II. Dodatky, doplòky a opravy biografických hesel „Seznamu biografií ès. Entomologù (entomologové nežijící I“ ve svazcích 1–15 z let 1979–1995. [A List of biographies of Czechoslovak entomologists. II. Additions and corrections to volumes 1–15 published in 1979–1995.] Klapalekiana 34(Suppl.): 1–238 (in Czech). KULA E. 2002: The leafhopper fauna in birch (Betula pendula Roth) stands. Journal of Forest Science 48: 351–360. KUNZ G., NICKEL H. & NIEDRINGHAUS R. 2011: Fotoatlas der Zikaden Deutschlands. Photographic atlas of the planthoppers and leafhoppers of Germany. WABV Fründ, Scheessel 293 pp. LAUTERER P. 1958: Pøíspìvek k poznání køísù ÈSR (Hom., Auchenorrhyncha) II. A contribution to the knowledge of the leaf-hoppers of Czechoslovakia (Hom., Auchenorrhyncha) II. Acta Musei Moraviae, Scientiae naturales (Brno) 43: 125–136 (in Czech, English summary). LAUTERER P. 1983: Fagocyba cerricola sp. n. and new and interesting records of leafhoppers from Czechoslovakia (Homoptera, Auchenorrhyncha). Acta Musei Moraviae, Scientiae naturales (Brno) 68: 139–152. LAUTERER P. 1995: Auchenorrhyncha. Pp. 165–175. In: ROZKOŠNÝ R. & VAÒHARA J.: Terrestrial Invertebrates of the Pálava Biosphere Reserve of UNESCO, I. Folia facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis, Biologia 92: 1–206. LAUTERER P. & N OVOTNÝ V. 1991: New findings of leafhoppers (Homoptera, Auchenorrhyncha) in Czechoslovakia. Acta Musei Moraviae, Scientiae Naturales 76: 265–268. LINNAVUORI R. 1969: Hemiptera IV. Animalia Fennica 13: 1–312. MALENOVSKÝ I. 2006: Køísi (Auchenorrhyncha, Hemiptera) CHKO Kokoøínsko. (Planthoppers and leafhoppers (Auchenorrhyncha, Hemiptera) of Kokoøínsko Protected Landscape Area.) Bohemia Centralis 27: 295–322 (in Czech, English abstract). MALENOVSKÝ I. 2013: RNDr. Pavel Lauterer, on the occasion of his eightieth birthday. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98: 000–000. MALENOVSKÝ I., BAÒAØ P. & K MENT P. 2011: A contribution to the faunistics of the Hemiptera (Cicadomorpha, Fulgoromorpha, Heteroptera, and Psylloidea) associated with dry grassland sites in southern Moravia (Czech Republic). Acta Musei Moraviae, Scientiae Biologicae 96: 41–187. MALENOVSKÝ I., BÜCKLE C., GUGLIELMINO A., KOCZOR S., NICKEL H., SELJAK G., SCHUCH S. & WITSACK W. 2013: Contribution to the Auchenorrhyncha fauna of the Pálava Protected Landscape Area (Czech Republic). Cicadina 13 (in press).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 261 I. MALENOVSKÝ

MALENOVSKÝ I. & BURCKHARDT P. 2003: Pavel Lauterer at Seventy. Acta Musei Moraviae, Scientiae biologicae 88: 181–190. MALENOVSKÝ I. & LAUTERER P. 2005a: Auchenorrhyncha (køísi). Pp. 147–155. In: FARKAÈ J., KRÁL D. & ŠKORPÍK M. (eds.): Èervený seznam ohrožených druhù Èeské republiky. Bezobratlí. Red list of threatened species in the Czech Republic. Invertebrates. Agentura ochrany pøírody a krajiny ÈR, Praha, 758 pp (in Czech and English). MALENOVSKÝ I. & LAUTERER P. 2005b: Leafhoppers, planthoppers and psyllids (Hemiptera: Cicadomorpha, Fulgoromorpha, Psylloidea) in ruderal habitats: material attracted by light in the suburbs of Brno (Czech Republic). Acta Musei Moraviae, Scientiae biologicae (Brno) 90: 195–207. MALENOVSKÝ I. & LAUTERER P. 2006: Auchenorrhyncha – Køísi. Pp. 266–271. In: MLÍKOVSKÝ J. & STÝBLO P. (eds.): Nepùvodní druhy fauny a flóry Èeské republiky. [Alien species of the fauna and flora of the Czech Republic]. Èeský svaz ochráncù pøírody, Praha, 496 pp (in Czech). MALENOVSKÝ I. & LAUTERER P. 2010: Additions to the fauna of planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) of the Czech Republic. Acta Musei Moraviae, Scientiae Biologicae 95: 49–122. MALENOVSKÝ I. & LAUTERER P. 2012: Leafhoppers and planthoppers (Hemiptera: Auchenorrhyncha) of the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). In: MALENOVSKÝ I., KMENT P. & K ONVIÈKA O. (eds.): Species inventories of selected insect groups in the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). Acta Musei Moraviae, Scientiae Biologicae 96(2) (2011): 155–322. MALENOVSKÝ I. & TROPEK R. 2009: Faunistic records from the Czech Republic – 274. Hemiptera: Cicadomorpha: Cicadellidae. Klapalekiana 45: 80–82. MELICHAR L. 1899: Beitrag zur Kenntniss der Homopteren-Fauna von Tunis. Wiener Entomologische Zeitung 18: 175–190. MEYER-ARNDT S. & REMANE R. 1992: Phylogenie und Speziation der Wagner, 1951 (Homoptera: Auchenorrhyncha: Cicadellidae). Teil 1. Marburger Entomologische Publikationen 2: 1–387. NAST J. 1972: Palaearctic Auchenorrhyncha (Homoptera). An annotated check list. Polish Scientific Publishers, Warszawa, 550 pp. NICKEL H. 2003: The Leafhoppers and planthoppers of Germany (Hemiptera, Auchenorrhyncha): Patterns and strategies in a highly diverse group of phytophagous insects. Pensoft Publishers, Sofia – Moscow and Goecke & Evers, Keltern, 460 pp. NIEDRINGHAUS R., BIEDERMANN R. & NICKEL H. 2010: Verbreitung der Zikaden des Großherzogtums Luxemburg – Textband. Ferrantia 60: 1–105. OSSIANNILSSON F. 1978: The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark. Part 1: Introduction, infraorder Fulgoromorpha. Fauna Entomologica Scandinavica Vol. 7(1). Scandinavian Science Press, Klampenborg, pp. 1–222. OSSIANNILSSON F. 1981: The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark. Part 2: The families Cicadidae, , Membracidae, and Cicadellidae (excl. Deltocephalinae). Fauna Entomologica Scandinavica, Vol. 7(2). Scandinavian Science Press, Klampenborg, pp. 223–593. OSSIANNILSSON F. 1983: The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark. Part 3: The family Cicadellidae: Deltocephalinae, Catalogue, Literature and Index. Fauna Entomologica Scandinavica Vol. 7(3). Scandinavian Science Press, Kopenhagen, pp. 594–979. PREYSSLER J. D. E. 1792: Beschreibungen und Abbildungen derjenigen Insekten, welche in Sammlungen nicht aufzubewahren sind, dann aller, die noch ganz neu, und solcher, von denen wir noch keine oder doch sehr schlechte Abbildung besitzen. Sammlung Physikalischer Aufsätze (Dresden) 2: 1–46, 1 tab. PREYSSLER J. D. E., LINDACKER J. T. & HOFER J. K. 1793: Beobachtungen über Gegenstände der Natur, auf einer Reise durch den Böhmerwald in Sommer 1791. Sammlung Physikalischer Aufsätze (Dresden) 3: 135–378. PRUNER L. & MÍKA P. 1996: Seznam obcí a jejich èástí v Èeské republice s èísly mapových polí pro síťové mapování fauny. List of settlements in the Czech Republic with associated map field codes for faunistic grid mapping system. Klapalekiana 32(Suppl.): 1–175. REMANE R. 1987: Zum Artenbestand der Zikaden (Homoptera: Auchenorrhyncha) auf dem Mainzer Sand. Mainzer Naturwissenschaftliches Archiv 25: 273–349. REMANE R. & FRÖHLICH W. 1994: Beiträge zur Chorologie einiger Zikaden-Arten (Homoptera Auchenorrhyncha) in der Westpaläarktis. Marburger Entomologische Publikationen 2(8): 131–188.

262 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New records of Auchenorrhyncha for Czech Republic

REMANE R. & GIUSTINA W. DELLA 1991: La Faune de France des Delphacidae (Homoptera Auchenorrhyncha). I. Récoltes d’août 1989. Cahiers des Naturalistes, Bulletin des Naturalistes Parisiens 47(2): 33–44. SÖDERMAN G. 2007. Taxonomy, distribution, biology and conservation status of Finnish Auchenorrhyncha (Hemiptera: Fulgoromorpha et Cicadomorpha). The Finnish Environment 7: 1–101 (available online at www.environment.fi/publications). SPITZER K. & DANKS H. V. 2006: Insect biodiversity of boreal peat bogs. Annual Review of Entomology 51: 137–161. SPITZNER W. 1892: Beitrag zur Hemipteren-fauna Mährens. Verhandlungen des Naturforschenden Vereines in Brünn 30(1891): 3–34. STEWART A. J. A. 2002: Techniques for sampling Auchenorrhyncha in grasslands. In: HOLZINGER W. E. (ed.): Zikaden. Leafhoppers, planthoppers and cicadas (Insecta: Hemiptera: Auchenorrhyncha). Denisia 4: 491–512. STRÜBING H. 1978: Euscelis ohausi Wgn. 1939 and Euscelis singeri Wgn. 1951: separate species or not? Auchenorrhyncha Newsletter 1: 15. ŒWIERCZEWSKI D. & WALCZAK M. 2011a: New records of leafhoppers for Poland (Hemiptera: Cicadomorpha). Polish Journal of Entomology 80: 291–298. ŒWIERCZEWSKI D. & WALCZAK M. 2011b: New and rare Auchenorrhyncha species in the Polish fauna. In: Bericht über die 17. Tagung des Arbeitskreises Mitteleuropäische Zikaden vom 27.–29. August 2010 in Mikulov (Tschechische Republik). DGaaE-Nachrichten 25(1): 32–33. SZWEDO J. & GÊBICKI C. 1998: Cicadella lasiocarpae Ossiannilsson, 1981 (Homoptera: Cicadodea: Cicadellidae) new to Polish fauna. Annals of the Upper Silesian Museum (Entomology) 8–9: 93–98. TISHECHKIN D. YU. 1999: Review of species of the genus Hephathus Ribaut, 1952 (Homoptera: Cicadellidae: Macropsinae) from the territory of Russia with notes on other Palaearctic species of the genus. Russian Entomological Journal 8: 239–252. TISHECHKIN D. YU. 2000: K voprosu o taksonomicheskom statuse Cicadella lasiocarpae (Homoptera, Cicadellidae). (On taxonomic status of Cicadella lasiocarpae (Homoptera, Cicadellidae)). Zoologicheskii Zhurnal 79: 863–867. TROPEK R., CERNA I., STRAKA J., CIZEK O. & KONVICKA M. 2013: Is coal combustion the last chance for vanishing insects of inland drift sand dunes in Europe? Biological Conservation 162: 60–64. TROPEK R., KADLEC T., HEJDA M., KOCAREK P., SKUHROVEC J., MALENOVSKY I., VODKA S., SPITZER L., BANAR P. & KONVICKA M. 2012: Technical reclamations are wasting the conservation potential of post-mining sites. A case study of black coal spoil dumps. Ecological Engineering 43: 13–18. VIDANO C. & ARZONE A. 1985: Zyginidia pullula: distribuzione nel territorio e ciclo biologico. Redia 68: 135–150. WEBB M. D. & VIRAKTAMATH C. A. 2009: Annotated check-list, generic key and new species of Old World leafhoppers with nomenclatorial changes in the Deltocephalus group and other Deltocephalinae (Hemiptera, Auchenorrhyncha, Cicadellidae). Zootaxa 2163: 1–64.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 265–271, 2013

New data on some rare planthoppers and leafhoppers in Poland (Hemiptera: Auchenorrhyncha)

KRZYSZTOF MUSIK & ARTUR TASZAKOWSKI Department of Zoology, Faculty of Biology and Environmental Protection, University of Silesia, Bankowa 9, 40-007, Katowice, Poland; e-mail: [email protected], [email protected]

MUSIK K. & TASZAKOWSKI A. 2013: New data on some rare planthoppers and leafhoppers in Poland (Hemiptera: Auchenorrhyncha). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 265–271. – This contribution presents new distribution data on nine rarely-collected species of planthopper and leafhopper in Poland: Pentastiridius beieri (Wagner, 1970), Asiraca clavicornis (Fabricius, 1794), Kelisia praecox Haupt, 1935, Stenocranus fuscovittatus (Stål, 1858), Aphrophora major Uhler, 1896, Stictocephala bisonia Kopp et Yonke, 1977, Handianus flavovarius (Herrich-Schäffer, 1835), Metalimnus steini (Fieber, 1869), and Sonronius dahlbomi (Zetterstedt, 1840). Distributional and ecological data are provided for each species. Keywords. Fulgoromorpha, Cicadomorpha, faunistics, central Europe, Silesian Upland, the Bia³owie¿a Primeval Forest, the Eastern Beskidy Mountains

Introduction The Polish fauna of Auchenorrhyncha comprises 538 species (CHUDZICKA 2004; £ABANOWSKI & SOIKA 1997; GÊBICKI 2003; ŒWIERCZEWSKI & GÊBICKI 2003; GAJ et al. 2009; ŒWIERCZEWSKI & STROIÑSKI 2011a, b; ŒWIERCZEWSKI & WALCZAK 2011; WALCZAK et al. 2012; MUSIK et al. 2013). This group of insects still requires a long-term field survey in Poland, since many species are known from only a few localities or have not been recorded for a long time; their actual distribution around the country is not clear. Further, many of the zoogeographical regions of Poland still lack proper study, and many new species for the country may thus await discovery. At present, faunistic studies are carried out mainly in the Silesian Upland and the Krakowsko-Wieluñska Upland. This contribution presents new localities for nine rarely-collected species of Auchenorrhyncha, found by the authors in the course of faunistic studies in the Silesian Upland, the Bia³owie¿a Primeval Forest, and the Eastern Beskidy Mountains.

Material and methods The material was collected in 2010–2013 with a standard sweep net and an aspirator. The insects were identified in the laboratory on the basis of external morphological characters and the anatomy of the male genitalia. Keys by BIEDERMAN & NIEDRINGHAUS (2009) and DLABOLA (1954) were used for identification. Ecological and biological data are taken from NICKEL (2003). The zoogeographical division of Poland adopted in this paper follows that presented in the Catalogue of Polish Fauna (NAST 1976). Codes of the fields of the UTM map grid are given in square brackets in the lists of material examined.

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All the material is held in the entomological collection of the Department of Zoology, University of Silesia, Katowice.

Results Family CIXIIDAE

Pentastiridius beieri (Wagner, 1970) Material examined. Silesian Upland, Ruda Œl¹ska, vicinity of Bieszowice II spoil tip [UTM:CA47], 06.vii.2012, 1 ♂ 2 ♀♀, leg. K. Musik. Remarks. The species is known from Austria, Germany, Italy, Switzerland, and Ukraine (NAST 1987, NICKEL 2003). Rarely collected in Poland, this planthopper is known from the western Beskids: Rabka-S³one and Myœlenice (NAST 1977) and the Silesian Upland: Rybnik, Ruda Œl¹ska and Bytom, where it was collected in the first half of the 1980s (ŒWIERCZEWSKI & GRUCA 2010). The new locality lies in Ruda Œl¹ska Bielszowice at the bottom of the Bielszowice II spoil tip. It is a sandy, dry site with scattered grasses as its main growth. According to the literature, P. beieri feeds on Salix spp. but it has also been collected from German tamarisk Myricaria germanica and Alder Alnus spp., and is considered stenotopic on the gravel banks of mountain rivers (NICKEL 2003).

Family DELPHACIDAE

Asiraca clavicornis (Fabricius, 1794)

Material examined. Silesian Upland, Piekary Œl¹skie Do³ki, spoil tip [UTM:CA57], 02.ix.2011, 1 ♀, leg. K. Musik. Remarks. This species is widely distributed in central and southern Europe, the Near East and North Africa (NAST 1972, 1987). In Poland, it has been recorded in the Pomeranian Lake District (WAGNER 1941), the Wielkopolsko-Kujawska Lowland (SZULCZEWSKI 1933), the Silesian Upland (SZULCZEWSKI 1931), the Krakowsko- Wieluñska Upland (SMRECZYÑSKI 1954), the Ma³opolska Upland, the Lubelska Upland (NAST 1976), and the Western Beskids (SMRECZYÑSKI 1906). Localities for A. clavicornis are scattered, mainly in the southern part of the country. The locality in Piekary Œl¹skie is a spoil tip covered in various species of grass and shrub. This species is usually found in dry, disturbed sites (NICKEL 2003).

Kelisia praecox Haupt, 1935 Material examined. Silesian Upland, Katowice Panweniki, Œlepiotka river valley [UTM:CA56], 30.v.2012, 1 ♀; 19.ix.2013, 3 ♂♂, leg. K. Musik. Remarks. In Europe this species has been recorded in Bulgaria, Czech Republic, Estonia, Germany, Greece, Latvia, Lithuania, Slovakia, and Yugoslavia (NAST 1987).

266 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New data on planthoppers and leafhoppers in Poland

Further distribution includes Iran (MOZAFFARIAN & WILSON 2011), Georgia, Kazakhstan, Russia, and Mongolia (NAST 1976). In Poland, K. praecox has been collected in the Krakowsko-Wieluñska Upland, the Western Beskidy Mountains (SMRECZYÑSKI 1954), and the the Sandomierska Lowland (BEDNARCZYK & GÊBICKI 1998). The locality cited here is the first in the Silesian Upland. It is a moist meadow near the Œlepiotka river and a forest edge. The species is frequently associated with Carex brizoides in moist sites, usually forest meadows or clearings (NICKEL 2003).

Stenocranus fuscovittatus (Stål, 1858) Material examined. Silesian Upland, Katowice Panweniki, Œlepiotka river valley [UTM:CA56], 19.iv.2013, 4 ♂♂, leg. K. Musik. Remarks. This Eurosiberian species is widely distributed in western, central, and eastern Europe including the Balkans (NAST 1972, 1987; BORODIN 2004), Georgia, Russia, Kazakhstan, China, and Mongolia (DLABOLA 1967, NAST 1972). In Poland, it is widely distributed and locally fairly common but rarely recorded. It feeds on certain species of Carex (C. elata, C. paniculata, C. panicea and others). It inhabits moist sites such as meadows, fens or spring mires (NICKEL 2003).

Family

Aphrophora major Uhler, 1896

Material examined. Eastern Beskidy Mountains, Libusza, disturbed meadow [UTM:EA10], 14.ix.2012, 1 ♀, leg. A. Taszakowski. Remarks. This Eurosiberian species has been recorded to date in Austria, Great Britain, Czech Republic, France, Germany, Italy, Switzerland, Netherlands, Ukraine and Yugoslavia (HOCH 2013). Further distribution includes central and northern Russia, Mongolia, north-eastern China, Japan and the Kuril Islands (ŒWIERCZEWSKI & GÊBICKI 2003). In Poland, it has been collected in the Masurian Lake District (ANDRZEJEWSKA 1965, GÊBICKI et al. 1982), the Bia³owie¿a Primeval Forest (NAST 1936), and the Lubelska Upland (BEDNARCZYK & GÊBICKI 1998). The last-cited locality is the southernmost record in Poland. A single female was collected in a disturbed meadow with shrubs. A. major is listed in the Polish Red Data Book as a species at lower risk (SZWEDO 2004); it is not protected by law. It feeds on birch Betula spp. and Salix spp. Its nymphal host plants are poorly known, but development probably takes place on Phragmites, Pedicularis, Salix and Artemisia (NICKEL 2003).

Family MEMBRACIDAE

Stictocephala bisonia Kopp et Yonke, 1977

Material examined. Eastern Beskidy Mountains, Libusza, disturbed meadow [UTM:EA10], 14.ix.2012, 1 ♂, leg. A. Taszakowski.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 267 K. MUSIK & A. TASZAKOWSKI

Remarks. Originally a Nearctic species introduced into Europe and currently present in Albania, Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, France, Germany, Greece, Hungary, Italy, Macedonia, Moldavia, Portugal, Romania, Slovakia, Slovenia, Spain, Switzerland, Ukraine and Yugoslavia (ARZONE et al. 1987, JANSKÝ et al. 1988, HOCH 2013). Recently recorded in Poland from localities near Warsaw, Rzeszów and Zagorzyce. This is the first record in Polish mountains. S. bisonia is a polyphagous species, with nymphs developing on a variety of herbaceous plants. Oviposition takes place on woody species (ŒWIERCZEWSKI & STROIÑSKI 2011a).

Family CICADELLIDAE

Handianus flavovarius (Herrich-Schäffer, 1835) Material examined. Eastern Beskidy Mountains, Wysowa-Zdrój, wet meadow [UTM:EV17], 07.viii.2012, 1 ♂ 2 ♀♀; Libusza, disturbed meadow [UTM:EA10], 14.ix.2012, 5 ♀♀, leg. A. Taszakowski. Remarks. An Eurosiberian species known from Czech Republic, Estonia, Germany, Latvia, Lithuania, Moldavia, Romania, Russia, Slovakia, Ukraine, and Yugoslavia (NAST 1972, 1987). Quite rare in Poland: it has been published from only three localities in the Bieszczady Mountains (the Tarnawa Nature Reserve; SZWEDO et. al 1998), the Pieniny Mountains (Zielone Ska³ki range; NAST 1973), and the Biebrza river valley in the Podlasie (GÊBICKI et al. 1982). It occurs in dry meadows and probably feeds on herbaceous Fabaceae (Vicia, Trifolium, Lotus) (MALENOVSKÝ et al. 2011).

Metalimnus steini (Fieber, 1869)

Material examined. Silesian Upland, Piekary Œl¹skie Do³ki, spoil tip [UTM:CA57], 02.ix.2011, 4 ♀♀, leg. K. Musik; Bia³owie¿a Primeval Forest, old railway [UTM: FD84], 25.vii.2012, 1 ♂, leg. K. Musik. Remarks. A Palaearctic species recorded from Austria, Azerbaijan, Czech Republic, Germany, Hungary, Japan, Russia and Ukraine (NAST 1972, NICKEL 2003, MALENOVSKÝ &LAUTERER 2010). It has been only recently reported from Poland and was known to date from four localities: Silesian Upland – Jaworzno; Krakowsko-Wieluñska Upland – Czêstochowa (Góra Kamienio³om, Stradom) and Ma³opolska Upland – Murawy Dobromierskie Nature Reserve (ŒWIERCZEWSKI & WALCZAK 2011, ŒWIERCZEWSKI & STROIÑSKI 2011b). The new locality in the Silesian Upland lies in the Piekary Œl¹skie suburban district at the bottom of a spoil tip. The specimens were collected in a moist pit covered in various species of grasses. The other locality, in the Bia³owie¿a Forest, is the most northern in Poland. One male specimen was collected as part of a faunistic survey of an old railway line in the Bia³owie¿a. This locality has a ruderal character, with species of grasses and shrubs. Metalimnus steini feeds on hairy sedge Carex hirta and may be encountered in ruderal habitats and meadows (NICKEL 2003).

268 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New data on planthoppers and leafhoppers in Poland

Sonronius dahlbomi (Zetterstedt, 1840) Material examined. Eastern Beskidy Mountains, Wysowa-Zdrój, disturbed meadow [UTM:EV17], 20.vii.2010, 1 ♂, leg. A. Taszakowski. Remarks. Rarely recorded in Poland, this Holarctic species is known from Belgium, Finland, France, Germany, Great Britain, Italy, Latvia, Norway, Slovakia, Sweden, Switzerland, Ukraine and Yugoslavia. Further distribution includes Russia, Kazakhstan and North America (NAST 1972, 1976, 1987). In Poland it was previously known from two localities: Mazowiecka Lowland (NAST 1936) and the Pieniny Mountains (NAST 1976). This is the first report from the Beskidy Mountains. One male specimen was collected in disturbed grassland in the Lower Beskids. Epilobium angustifolium, , Alchemilla sp. and Filipendula ulmaria have been cited as host plants (NICKEL 2003).

Conclusions Recent field surveys carried out in the Silesian Upland, the Bia³owie¿a Primeval Forest and the eastern Beskidy Mountains have revealed a number of rarely-collected species of Auchenorrhyncha. Perhaps the most interesting species, Pentastiridius beieri, was collected at the bottom of a mining spoil tip in Ruda Œl¹ska. This locality is an example of how mining spoil tips can provide substitute habitats for rare and specialized species: according to the literature, P. beieri is a specialist in the gravel banks of mountain rivers (NICKEL 2003). Additional recent localities for the Nearctic treehopper Stictocephala bisonia may suggest that this species is currently already widespread in southern Poland and that the number of known localities may increase rapidly in the near future. The distribution of Metalimnus steini still needs to be assessed by a more detailed survey, but the locality in the Bia³owie¿a Primeval forest suggests that this species is widely distributed, at least in southern and north-eastern Poland. With an increasing number of field surveys focused on the Auchenorrhyncha in the different zoogeographical regions of Poland, one may presume that in the near future many additional interesting and rare species will be recorded, including new species for the country, as the current number of 538 species in Poland cannot be considered as definitive.

References

ANDRZEJEWSKA L. 1965: Stratification and its dynamics in meadow communities of Auchenorrhyncha (Homoptera). Ekologia Polska 13: 685–715. ARZONE A., VIDANO C. & ALMA A. 1987: Auchenorrhyncha introduced into Europe from the Nearctic region: taxonomic and phytopathological problems. Pp. 3–17. In: WILSON M. R. & NAULT L. R. (eds.): Proceedings of 2nd International Workshop on Leafhoppers and Planthoppers of Economic Importance, Provo, Utah, USA, 28th July-1st Aug. 1986, CIE. London, xiii + 368 pp.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 269 K. MUSIK & A. TASZAKOWSKI

BEDNARCZYK J. & GÊBICKI C. 1998: Piewiki (Homoptera, Auchenorrhyncha) okolic Janowa Lubelskiego. Fragmenta Faunistica 41: 233–245. BIEDERMANN R. & NIEDRINGHAUS R. 2009: The Plant- and Leafhoppers of Germany. Identification key to all species. WABV Fründ, Scheeßel, 409 pp. BORODIN O. 2004: A checklist of the Auchenorrhyncha of Belarus (Hemiptera, Fulgoromorpha et Cicadomorpha). Beiträge zur Zikadenkunde 7: 29–47. CHUDZICKA E. 2004: Piewiki (Auchenorrhyncha = Cicadomorpha + Fulgoromorpha). [Leafhoppers and planthoppers.] Pp. 178–192. In: BOGDANOWICZ W., CHUDZICKA E., PILIPIUK I. & SKIBIÑSKA E. (eds.): Fauna Polski – charakterystyka i wykaz gatunków. Tom I. (Fauna of Poland – characterisation and checklist of species.) Muzeum i Instytut Zoologii PAN, Warszawa, 509 pp (in Polish, English summary). DLABOLA J. 1954: Køísi - Homoptera. Fauna ÈSR 1. [Leafhoppers - Homoptera. Fauna of the ÈSR 1.] Nakladatelství Èeskoslovenské akademie vìd, Praha, 399 pp (in Czech, German and Russian summary). DLABOLA J. 1967: Ergebnisse der zoologischen Forschungen von Dr Z. Kaszab in der Mongolei. Acta Faunistica Entomologica Musei Nationalis Pragae 12: 137–152. GAJ D., WALCZAK M. & WOJCIECHOWSKI W. 2009: Communities of planthoppers and leafhoppers (Hemiptera: Fulgoromorpha et Cicadomorpha) inhabiting selected plant communities in the Sto³owe Mountains National Park. Annals of the Upper Silesian Museum in Bytom, Natural History, Monograph 19: 1–174. GÊBICKI C. 2003: Nowe stanowisko Utecha lugens (Germar, 1821) w Polsce (Hemiptera, Ulopidae). (New locality of Utecha lugens (German, 1821) in Poland (Hemiptera: Ulopidae).) Acta Entomologica Silesiana 9-10: 23–26 (in Polish, English abstract). GÊBICKI C., BARTNICKA I., BOK£AK E. & MA£KOWSKI E. 1982: Piewiki (Homoptera, Auchenorrhyncha) Kotliny Biebrzy. (Leafhoppers (Homoptera, Auchenorrhyncha) of the Biebrua Valley.) Acta Biologica (Katowice) 10: 13–21 (in Polish, English abstract). HOCH H. 2013: Fauna Europaea: Hemiptera: Cicadomorpha etc. Fauna Europaea, version 2.6.1. Available online at http://www.faunaeur.org (last accessed on 27 August 2013). JANSKÝ V., KRIŠTÍN A. & OKÁLI I. 1988: Der gegenwärtige Stand der Verbreitung und neue Erkenntnisse über die Bionomie der Art Stictocephala bisonia (Homoptera, Membracidae) in der Slowakei. Biológia (Bratislava) 43: 527–533. £ABANOWSKI G. & SOIKA G. 1997: Nowe i mniej znane szkodniki wystêpuj¹ce na drzewach i krzewach ozdobnych. [New and little known pests ocurrïng on ornamental trees and shrubs.] Progress in Plant Protection 31: 218–223 (in Polish). MALENOVSKÝ I., BAÒAØ P. & KMENT P. 2011: A contribution to the faunistics of the Hemiptera (Cicadomorpha, Fulgoromorpha, Heteroptera, Psylloidea) associated with dry grassland sites in southern Moravia (Czech Republic). Acta Musei Moraviae, Scientiae biologicae (Brno) 96(1): 41–187. MALENOVSKÝ I. & LAUTERER P. 2010: Additions to the fauna of planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) of the Czech Republic. Acta Musei Moraviae, Scientiae Biologicae (Brno) 95: 49–122. MOZAFFARIAN F. & WILSON M. R. 2011: An annotated checklist of the planthoppers of Iran (Hemiptera, Auchenorrhyncha, Fulgoromorpha) with distribution data. ZooKeys 145: 1–57. MUSIK K., WALCZAK M., DEPA £., JUNKIERT £. & JEDYNOWICZ A. 2013: Trigonocranus emmeae Fieber, 1876 (Hemiptera, Fulgoromorpha, Cixiidae) – a new species for Poland. In: POPOV A., GROZEVA S., SIMOV N. & TASHEVA E. (eds.): Advances in Hemipterology. ZooKeys 319: 249–253. NAST J. 1936: Nowe dla Polski lub mniej znane gatunki Homoptera. (Neue für Polen oder weniger bekannte Homopteren-Arten.) Fragmenta Faunistica (Warszawa) 2: 323–326 (in Polish, German summary). NAST J. 1972: Palearctic Auchenorrhyncha (Homoptera). An annotated check list. Polskie Wydawnictwo Naukowe, Warszawa, 550 pp. NAST J. 1973: Uzupe³nienia i sprostowania do fauny Auchenorrhyncha (Homoptera) Polski. Fragmenta Faunistica (Warszawa) 19: 39–53 (in Polish). NAST J. 1976: Piewiki. Auchenorrhyncha (Cicadodea). Catalogus faunae Poloniae 25, XXI (1). Polskie Wydawnictwo Naukowe, Warszawa, 256 pp (in Polish). NAST J. 1977: Homopterological notes XIII-XX. Annales Zoologici (Warszawa) 34: 27–37. NAST J. 1987: The Auchenorrhyncha (Homoptera) of Europe. Annales Zoologici (Warszawa) 40: 535–661. NICKEL H. 2003: The Leafhoppers and Planthoppers of Germany (Hemiptera, Auchenorrhyncha): Patterns and strategies in a highly diverse group of phytophagous insects. Series Faunistica 28. Pensoft Publishers, Sofia – Moscow, Goecke & Evers, Keltern, 460 pp.

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SMRECZYÑSKI S. 1906: Wykaz pluskwiaków nowych dla fauny galicyjskiej. [List of new Hemiptera for the fauna of Galicia.] Sprawozdania Komitetu Fizjograficznego (Kraków) 40: 72–79. SMRECZYÑSKI S. 1954: Materia³y do fauny pluskwiaków (Hemiptera) Polski. [Materials to the Hemiptera fauna of Poland.]. Fragmenta Faunisctica (Warszawa) 7: 1–146. SZULCZEWSKI J. W. 1931: Notatki entomologiczne i zoocedidiologiczne z powiatu lublinieckiego na Górnym Œl¹sku. [Entomological and zoocecidological notes from the environs of Lublin in Upper Silesia.] Polskie Pismo Entomologiczne 10:124–141. SZULCZEWSKI J. W. 1933: Beitrag zur Cicadenfauna des Posener Landes. Deutsche Wissenschaftliche Zeitschrift für Polen (Poznañ) 26: 95–103. SZWEDO J. 2004: Aphrophora major Uhler, 1896. Piewiki Hemiptera. ‘Auchenorrhyncha’, Fulgoromorpha et Cicadomorpha. Pp. 79–80. In: G£OWACIÑSKI Z. & NOWACKI J. (eds.): Polska Czerwona Lista Zwierz¹t, II. Bezkrêgowce. [Polish Red Book of Animals, II. Invertebrates]. Instytut Ochrony Przyrody PAN, Kraków, 447 pp (in Polish). SZWEDO J., GÊBICKI C. & WÊGIEREK P. 1998: Leafhopper communities (Homoptera, Auchenorrhyncha) of selected peat-bogs in Poland. Annals of the Upper Silesian Museum, Natural History 15: 154–176. ŒWIERCZEWSKI D. & GÊBICKI C. 2003: Ró¿norodnoœæ gatunkowa piewików w Polsce i jej ochrona (Hemiptera, Auchenorrhyncha). [Diversity of species of Hemiptera in Poland and their protection (Hemiptera, Auchenorrhyncha).] Acta Entomologia Silesiana 9-10: 77–84 (in Polish, English abstract). ŒWIERCZEWSKI D. & GRUCA P. 2010: Rare leafhopper species in Polish fauna – distributional maps (Hemiptera: Fulgoromorpha et Cicadomorpha). Chemistry, Environment, Biotechnology 1: 41–99. ŒWIERCZEWSKI D. & STROIÑSKI A. 2011a: The first records of the Nearctic treehopper Stictocephala bisonia in Poland (Hemiptera: Cicadomorpha: Mambracidae) with some comments on this potential pest. Polish Journal of Entomology 80: 13–22. ŒWIERCZEWSKI D. & STROIÑSKI A. 2011b: Planthoppers and leafhoppers of the Przedborski Landscape Park (Hemiptera: Fulgoromorpha et Cicadomorpha). Polish Journal of Entomology 80: 277–290. ŒWIERCZEWSKI D. & WALCZAK M. 2011: New records of leafhoppers for Poland (Hemiptera: Cicadomorpha). Polish Journal of Entomology 80: 291–298. WAGNER W. 1941: Die Zikaden der Provinz Pommern. Dohrniana 20: 95–184. WALCZAK M., MUSIK K. & MOKRZYCKA A. 2012: (Osborn, 1900) – a new leafhopper for Polish fauna (Hemiptera: Cicadomorpha). Wiadomoœci Entomologiczne 31: 242–250.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 273–315, 2013

Zur Etymologie der Zikadennamen Mittel- und Nordeuropas [On the etymology of Auchenorrhyncha names of central and northern Europe]

HERBERT NICKEL Ehrengard-Schramm-Weg 2, 37085 Göttingen, Germany; e-mail: [email protected]

NICKEL H. 2013: Zur Etymologie der Zikadennamen Mittel- und Nordeuropas. [On the etymology of Auchenorrhyncha names of central and northern Europe.] In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 273–315. – This paper gives German translations of scientific names of all currently recognized genera, subgenera, species and subspecies of the central and north European fauna of planthoppers and leafhoppers (Hemiptera: Auchenorrhyncha) including some synonyms. Altogether 363 supraspecific and 946 specific and subspecific names are treated. Remarks on the interpretation are given in some non-obvious cases. Keywords. Hemiptera, Auchenorrhyncha, Fulgoromorpha, Cicadomorpha, etymology

Einleitung Die vorliegende Arbeit soll einen Beitrag zum sprachlichen Verständnis der wissenschaftlichen Namen der Zikaden darstellen, wie er bereits für zahlreiche Insektengruppen vorliegt, z.B. für die Coleoptera (SCHENKLING 1917), Heteroptera (nur Gattungsnamen, s. REDL & KALLENBORN 1995), Odonata (FLIEDNER 1997) und Rhaphidioptera (ASPÖCK & ASPÖCK 2013). Behandelt werden alle Arten, Unterarten, Gattungen und Untergattungen sowie wichtige Synonyme aus Mittel- und Nordeuropa, einschließlich Weißrussland, Mittelrussland, Polen, der Slowakei, Ungarn, Slowenien, Norditalien und der Nordhälfte Frankreichs. Die Auswahl erfolgte durch eine Filterung der Datenbank des Autors mit den Zikadenarten Europas, welche im Wesentlichen auf NAST (1987) basiert und seitdem anhand neuer Literatur ergänzt wird (DEN BIEMAN et al. 2011, BORODIN 2004, D’URSO 1995, GÜNTHART & MÜHLETHALER 2002, GYÖRFFY et al. 2009, HOLZINGER 2009, HOLZINGER & SELJAK 2001, MÜHLETHALER et al. 2009, SÖDERMAN et al. 2009, NICKEL 2010 sowie zahlreiche Einzelarbeiten verschiedener Autoren). Die Gattungsnamen der Zikaden sind in der Regel – wie allgemein in der zoologischen Nomenklatur üblich – aus dem Griechischen abgeleitet (wenn auch häufig durch die Endung latinisiert). Allerdings gibt es hiervon auch Ausnahmen: Henri Ribaut hat sich aus dem Baskischen bedient, William Distant aus dem Sanskrit, Shonen Matsumura und Tamotsu Ishihara aus dem Japanischen. Seit dem 20. Jahrhundert schaffen besonders amerikanische Autoren (Bryan Beirne, Merv Nielson, Paul Oman u.a.) Kunstnamen, die akronymisch aus verschiedenen Wörtern, z.T. auch Namen zusammengesetzt sind, welche ohne Angabe der derivatio nominis oder ohne persönlichen Kontakt mit dem Autor so gut wie gar nicht mehr deutbar sind. Als Sonderfall soll hier noch eine Reihe von zunächst gar nicht erklärbaren Namen, v.a. aus

273 H. NICKEL dem Zeitraum vor 1860, erwähnt werden. Diese gehen auf eine scheinbar wahllose Benennung nach Städten und anderen Orten bzw. deren Bewohnern aus der griechischen Antike zurück, z.B. Issus Fabricius, 1803 und Iassus Fabricius, 1803 (KERZHNER 2007), außerdem auch Oliarus Stål, Ledra Fabricius, 1803 und Myndus Stål, 1862. Die Artnamen stammen mit nur wenigen Ausnahmen (z.B. von Alexej Zachvatkin) aus dem Lateinischen. Neben charakteristischen morphologischen Merkmalen oder Färbungselementen werden häufig auch geografische Namen (z.B. der locus typicus oder auch Regionen) benannt und Fachkollegen, Familienmitglieder und Freunde geehrt. Wie bei vielen anderen phytophagen Insekten wurden auch die Wirtspflanzen oder auch die Lebensräume des Öfteren zur Benennung herangezogen. Willkürlich zusammengesetzte Namen sind hier bei den Artnamen (zumindest in Europa) selten, kommen aber zunehmend in Mode (z.B. Christoph Bückle, Adalgisa Guglielmino, Reinhard Remane). Je nach linguistischer bzw. philologischer Affinität des jeweiligen Beschreibers wurden die Namen mit sprachlich unterschiedlicher Korrektheit gebildet. Besonders bei der Formung von Flexionsendungen, Suffixen und Präfixen unterlaufen den Autoren gelegentlich Fehler, welche dann die richtige Deutung des gesamten Namens erschweren können. Die Quellenlage ist heute sicherlich recht komfortabel, was v.a. an der digitalen Verfügbarkeit vieler Wörterbücher liegt. Unter den benutzten Wörterbüchern sind insbesondere WERNER (1968), GEMOLL (1997), STOWASSER et al. (1979) und BROWN (1954) zu nennen, außerdem HENTSCHEL & WAGNER (1996), GENAUST (1996), Wikipedia und andere Online-Quellen (ANONYMUS 2013b, c, d). Schließlich wurde der Pollux Archimedes Project Dictionary Access benutzt (ANONYMUS 2013a), ein Online- Wörterbuch, das Zugriff auf LIDELL, SCOTT & JONES (A Greek–English Lexicon), LEWIS & SHORT (Latin Dictionary) und weitere Quellen gewährt. Trotz intensiver Recherche einschließlich der Sichtung zahlreicher Originalbeschreibungen und umfangreicher Korrespondenz mit Kollegen konnten nicht alle Namen ausreichend geklärt werden, und auch Irrtümer und Fehlübersetzungen bzw. – deutungen sind nicht ausgeschlossen. Der Verfasser ist für alle diesbezüglichen Hinweise dankbar. Die Erstbeschreiber werden hier sowohl für die Arten als auch für die Gattungen abgekürzt mitaufgeführt. Bei Artnamen, welche in mehreren Gattungen auftreten, wird nur ein einziger (i.d.R. der ältere) Beschreiber aufgeführt, von dem auch das grammatikalische Genus übernommen wurde.

Verwendete Abkürzungen:

bask. = baskisch lat. = lateinisch engl. = englisch nlat. = neulateinisch franz. = französisch pers. = persisch griech. = griechisch russ. = russisch hebr. = hebräisch sanskr. = sanskrit japan. = japanisch span. = spanisch kasach. = kasachisch tatar. = tatarisch

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Liste der Gattungen und Untergattungen Acanalonia Spin. mit unregelmäßigem roten Hof (griech.) Acanthocixius W. Wg. der mit Stachel versehene Cixius (griech.) – siehe dort Acanthodelphax Le Q. wörtlich: das Dornenferkel (griech.) Acericerus Dlab. frei: der mit Hörnern versehene auf dem Ahorn (lat./griech.) Achilida Stål vermutlich: die Lippenlose (griech.) Achorotile Fieb. von achoros (griech.: ohne Tanz, traurig); frei: die Freudlose Aconurella Rib. die mit spitzem Schwänzchen (lat.) Acucephalus Germ. der Spitzkopf (griech.) Adarrus Rib. frei: der mit Hörnern versehene (bask.: adar = Horn, Bogen), nach der Form der Penisanhänge Agallia Curt. die Jubelnde (griech.) Agalmatium Em. Puppe, Statuette (griech.) Aglena Am. et Serv. die ohne Augen (griech.) Aguriahana Dist. nach der indischen Volksgruppe der Aguriah, die als Eisenschmelzer bekannt waren; die Typusart A. metallica Dist. ist aus der selben Region beschrieben und hat einen metallischen Fleck im Vorderflügel Alebra Fieb. vermutlich: die Wohlgenährte (griech.) Allygus Fieb. an Weidenrute (von griech.: lygos) Alnetoidia Dlab. kombiniert aus Alnus (lat.: Erle) und Idia (siehe dort) Anaceratagallia Zachv. die von der (nordamerikanischen) Aceratagallia Kirk. verschiedene (griech.) Anakelisia W. Wg. im Sinne von Wagner: die über Kelisia (siehe dort) hinaus entwickelte (griech.) Anomia Fieb. die aus der Reihe fallende; wörtlich: die Unregelmäßige (griech.) Anoplotettix Rib. die waffenlose Zirpe (griech.) Anoscopus Kbm. der Emporschauende (griech.) Anoterostemma P. Löw von anoteros (griech.: höher, am höchsten), stemma (griech.: Kopfbinde, Kranz), bezogen auf die nach hinten gerückten Ozellen Apartus Hlz. akronymisch nach einem österreichischen Forschungs- förderprogramm benannt (Austrian Programm for Advanced Research and Technology) Aphrodes Curt. die Schaumartige (griech.), vermutlich bezogen auf eine (scheinbare) Ähnlichkeit mit Schaumzikaden Aphrophora Germ. Schaum tragend (griech.) Araeopus Spin. der schlanke Fuß (griech./lat.) Araldus Rib. von aralde (bask.: Gruppe, Menge, auch Paar), Bezug unklar Arboridia Zachv. frei: die eigenartige Baumbewohnerin (lat./griech.) Ariellus Rib. vermutlich: der kleine Widder (lat.) Arocephalus Rib. der Pflugköpfige (griech.)

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Arrailus Rib. von arrailu (bask.: Spalt, Ritze), bezogen auf die Aedeagus- form Arthaldeus Rib. frei: der zur Schafherde gehörende (bask.: artalde = Schaf- herde) Artianus Rib. vermutlich: der dazwischen stehende (bask.: arte = Zwischen- raum) Asiraca Latr. die Heuschrecke (griech.) Asymmetrasca Dlab. kombiniert aus asymmetria (griech.: Asymmetrie) und Empoasca (siehe dort) Athysanus Burm. der Ungefranste (griech.) Atropis Kbm. ohne Kiel (griech.) Aurkius Rib. von aurki (bask.: davor, gegenüber), vermutlich bezogen auf die Position des Gonoporus Austroasca Lower von austro (lat.: die südliche) und Empoasca (siehe dort) Balcanocerus M.-C. nach der Typusart Idiocerus balcanicus Horv.; frei: der mit Hörnern versehene vom Balkan (lat./griech.) Balclutha Kirk. nach einem gleichnamigen Dorf in Schottland benannt Bathysmatophorus J. Shlb. von bathysma (griech.: Tiefe) und phorós (griech.: tragend), eventuell bezogen auf den herabgedrückten Vorderkörper Batracomorphus Lew. der Froschförmige (griech.) Belaunus Rib. von belaun (bask.: Ellenbogen, Knie), bezogen auf den Knick im Aedeagusschaft Bobacella Kusn. nach dem Fundort des Holotypus in Bauen des Steppen- murmeltiers (Marmota bobak) Boreotettix Ldb. die Nordzirpe (lat./griech.) Brachyceps Kirk. der Kurzkopf (griech.) Bugraia Koçak benannt nach dem Sohn des Beschreibers, Bugra Kocak Bursinia A. Costa die Ledrige (griech.) Bythoscopus Germ. wörtlich: der Tiefenschauer (griech.), bezogen auf die nach unten gerichteten Ozellen Calamotettix Em. die (Schilf-)Rohrzikade (griech.) Caliscelis Lap. die mit schönen Beinen (griech.) Calligypona J. Shlb. kallos = Schönheit, gypos = krumm, gebogen (griech.), eventuell auch von gypona (griech.: Springerin) Centrotus F. der Bestachelte, von kentron (griech.: Stachel) Ceratocixius W. Wg. der gehörnte (griech.) Cixius (siehe dort) Cercopis F. von kerkops: der Spitzbube (griech.) Ceresa Am. et Serv. der Haken (hebr.) Chanithus Kol. der Stecher (von hebr. chanith = die Pike), bezogen auf die Kopfspitze Chelidinus Em. von chelidon (griech.: Schwalbe), bezogen auf das schwal- benschwanzartig gegabelte Konnektiv Chiasmus M. et R. die Überkreuzung (griech./nlat.), nach den sich über- lappenden Elytren

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Chloothea Em. die Grasgöttin (griech.) Chloriona Fieb. frei: die Begrünte (griech.) Chlorionidea P. Löw ähnlich Chloriona (griech.) Chlorita Fieb. die mit Grün versehene (griech.) Chloroneura Walsh frei: die mit grünen Flügeladern (griech.) Chunrocerus Zachv. abgeleitet von der nahestehenden Gattung Chunra Dist. (vermutlich sanskr., eventuell: Pfauenauge, bezogen auf die runden Flecken im Costalbereich) und kéras (griech.: Horn) Cicada L. die Zikade, Baumgrille (lat.) Cicadatra Kol. von Cicada (siehe dort) und atra (lat.: schwarz) Cicadella Latr. die kleine (Sing-) Zikade (lat.) Cicadetta Kol. die kleine Cicada (siehe dort) Cicadivetta Boul. von Cicadetta, mit neuer, willkürlicher Endung Cicadula Zett. die kleine Zikade (lat.) Circulifer Zachv. der Ringträger (lat.), bezogen auf die Penisform Cixidia Fieb. der eigenartige Cixius (siehe dort) Cixius Latr. eventuell von ciccus = kleine Heuschrecke (lat.) Colladonus Ball vermutlich von collum (lat.: Hals) und donum (lat.: Geschenk) Colobotettix Rib. die verstümmelte Zirpe (griech.), bezogen auf die Penisform Conomelus Fieb. von konos (griech.: Kegel) und melos (griech.: Glied), bezogen auf das zweite Antennenglied Conosanus Osb. et Ball vermutlich: Kegelafter (griech./lat.), nach der Form des Genitalsegmentes Coryphaelus Put. aus Umbenennung des präokkupierten Namens Coryphaeus Fieb. entstanden (griech.: kóryphaios = Anführer) Cosmotettix Rib. die geschmückte Zirpe (griech.) Coulinus Beirne eventuell Bezug zum Lebensraum: coulée (franz./engl. = Tal, Rinne) Criomorphus Curt. der Widderförmige (griech.) Cyperana De L. nach der Wirtspflanze Cyperus Cyphopterum Mel. von griech. kyphos (Buckel) und pteron (Flügel) Delphacinus Fieb. der dem Delphax ähnliche (siehe dort) Delphacodes Fieb. der dem Delphax ähnliche (siehe dort) Delphax F. das Ferkel (griech.), bezogen auf die entfernt Schweine- ohren ähnelnden abgeplatteten Antennengrundglieder Deltocephalus Burm. der dreieckige Kopf (griech.) Dicranotropis Fieb. die mit zwei Kielen auf dem Schädel versehene (griech.) Dictyophara Germ. frei: die ein Netzgewand trägt (griech.) Dikraneura Hd. die mit gegabelten Adern (griech.) Dimissalna Boul. von der Typusart dimissa (siehe dort), mit willkürlicher Endung Diplocolenus Rib. der mit doppelter Scheide versehene (griech.), bezogen auf die Penisform

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Ditropis Kbm. der doppelte Kiel (griech.) Ditropsis W. Wg. im Sinne des Autors vermutlich “der doppelte Kiel” (griech.), der korrekterweise allerdings Ditropis (siehe dort) heißen müsste, dieser Name war aber bereits vergeben Doliotettix Rib. die listige Zirpe (griech.), Bezug unklar Doratulina Mel. einer kleinen Doratura ähnlich (siehe dort) Doratura J. Shlb. der Spießschweif (griech.), bezogen auf die Form des Ovipositors Dorycephalus Kouch. der Lanzenköpfige (griech.) Dorydium Burm. von dory (griech.: Lanze) Dryocyba Vilb. zusammengezogen aus drys: (griech.: Baum) und Typhlocyba (siehe dort) Dryodurgades Zachv. von drys: (griech.: Baum) und durga (sanskr.), eine vielarmige Göttin im Hinduismus; vermutlich bezogen auf die Aedeagusanhänge Dudanus Dlab. nach dem tschechischen Hemipterologen Ladislav Duda (1854–1895) benannt Ebarrius Rib. eventuell von barri (bask.: neu) Ederranus Rib. von eder (bask.: hübsch) Edwardsiana Zachv. nach dem britischen Hemipterologen James Edwards (1856–1928) benannt Elymana De L. nach der in dieser Gattung beliebten Wirtspflanze Elymus Emelyanoviana An. nach dem russischen Zikadologen und Entomologen Alexandr Fedorovich Emelyanov (* 1936) benannt Empoasca Walsh von der amerikanischen Gattung Empoa Fitch abgeleitet, Bedeutung unklar, eventuell von (griech.: Stechmücke, Gnitze) Enantiocephalus Hpt. wörtlich: der gegenüberliegende Kopf (griech.), vermutlich bezogen auf den wie das Körperhinterende zugespitzten Kopf Endria Oman vermutlich nach einem weiblichen Vornamen, Bezug jedoch unklar Eohardya Zachv. vermutlich: die ursprüngliche Form (im evolutiven Sinn) von Hardya (siehe dort) Epiptera Metc. von epi (griech.: auf) und pteron (griech.: Flügel), vermutlich bezogen auf die übereinander liegenden Flügel Erasmoneura Young frei: die mit lieblichen Flügeladern (griech.) Erdianus Rib. vermutlich: der Mittlere (bask.) Erotettix Hpt. die Liebeszirpe (griech.) Errastunus Rib. von bask.: erraste = leicht, bequem; Bezug jedoch unklar Errhomenus Fieb. der Starke (griech.) Erythria Fieb. von erythros (griech.: rot) Erzaleus Rib. frei: der Uferbewohner (von bask. ertz = Ufer, Saum)

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Euconomelus Hpt. der schöne Conomelus (siehe dort) Euides Fieb. die Schöne (griech.) Eupelix Germ. der besondere Helm (griech.) Eupterycyba Dlab. die Eupteryx-köpfige (siehe dort) Eupteryx Curt. der besondere Flügel trägt Eurhadina Hpt. die besonders verschmälerte, schlanke (griech.) Eurybregma Scott breiter Vorderkopf (griech.) Eurysa Fieb. von eurys (griech.: breit) Eurysanoides Hlz., Kam. et Nick. die der Eurysa ähnliche (siehe dort) Eurysella Em. eine kleine Eurysa (siehe dort) Eurysula Vilb. eine kleine Eurysa (siehe dort) Euscelidius Rib. der Euscelis ähnliche Euscelis Br. der mit typischen Beinen (lat.) Evacanthus Le P. et S. der mit einem besonderen Dorn versehene (griech.) Exitianus Ball vermutlich von exitus (lat.: Tod, Ausgang), Bezug auf den langen Ovipositor, Bedeutung jedoch unklar Fagocyba Dlab. zusammengezogen aus Fagus (lat.: Buche) und Typhlocyba (siehe dort) Falcidius Stål von falx (lat.: Sichel) Falcotoya Fenn. kombiniert aus falx (lat.: Sichel) und Toya (siehe dort) Ficocyba Vid. kombiniert aus Ficus (lat.: Feige) und -cyba (siehe Tyhlocyba) Fieberiella Sign. nach dem böhmischen Hemipterologen Franz Xaver Fieber (1807–1872) benannt Flammigeroidea Dlab. ähnlich einer Flammenträgerin (lat.) Flastena Nast laut Nast eine willkürliche Buchstabenkombination Flata F. die Bläserin, Lautererzeugerin (lat.) Florodelphax Vilb. nach dem livländischen Hemipterologen Gustav August Adam von Flor (1829–1883) benannt, delphax (griech.) = Ferkel Forcipata De L. et Caldw. die mit einer Zange versehene (lat.) Fruticidia Zachv. kombiniert aus frutex (Genitiv: fruticis) = Strauch, und Idia (siehe unten); frei: die eigenartige Strauchbewohnerin (lat./griech.) – der manchmal verwendete Name Frutioidia basiert auf einem lapsus calami Fulgora L. römische Göttin des Blitzes, bezogen auf das der Typusart F. laternaria L. nachgesagte nächtliche Leuchten des Vorderkopfes Gargara Am. et Serv. von gargar (hebr.: Korn), bezogen auf die Körperform Glossocratus Fieb. wörtlich: der Zungenstarke (griech.) Goldeus Rib. von golde (bask.: Pflug), bezogen auf die Aedeagusform Goniagnathus Fieb. der mit gewinkelter Wange (griech.)

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Graphocephala Van Duz. mit gezeichnetem Kopf (griech.) Graphocraerus Thoms. vermutlich: der durch seine Zeichnung unterschiedene (griech.) Gravesteiniella W. Wg. nach dem niederländischen Hemipterologen Willem Hendrik Gravestein (1906–1989) benannt Grypotes Fieb. Krummnasigkeit (griech.), bezogen auf den in Seitenansicht gebogenen Frontoclypeus Gyponana Ball die Springerin (griech.) Haematoloma Hpt. die Blutrandige (griech.) Handianus Rib. von handi (bask.: groß) Hardya Edw. nach dem britischen Entomologen James Hardy (1815–1898) benannt Hauptidia Dwor. nach dem Hallenser Homopterologen Hermann Haupt (1873–1959) benannt Hecalus Stål der entspannt ruhende (griech.) Henschia Leth. nach dem slowakischen Arzt und Entomologen Andrija Hensch (1857–1930) benannt Hephathus Rib. der Dunkle (hebr.), nach einem Mononym von Amyot (1848) Hesium Rib. das Abgezäunte (bask.: hesi = Einfriedung) Hishimonoides Ishih. der dem Hishimonus ähnliche (nlat.; siehe dort) Hishimonus Ishih. von hishi (japan.: Raute) und mon (japan.: Zeichnung) Horvathianella An. nach dem ungarischen Hemipterologen Géza Horváth (1847–1937) benannt Hyalesthes Sign. das gläserne Kleid (griech.), nach den hyalinen Flügeln Hyledelphax Vilb. das Waldferkel (griech.) Hypericiella Dwor. von Hypericum (Johanniskraut) abgeleitet Hysteropterum das seiner Flügel beraubte (griech.) Am. et Serv. Iassus F. griech. Hafenstadt in Karien, heute Südwest-Türkei Idia Fieb. die Eigenartige (griech.) Idiocerus Lew. der mit besonderen Hörnern (= Antennen) versehene (griech.) Idiodonus Ball eventuell der eigenartig beschenkte (griech./lat.) Igutettix Mats. vermutlich nach dem Sammler des Holotypus, S. Iguchi, benannt Indiagallia Hlz. nach Indi (Kosename von Ingrid) benannt Irinula Rib. zur Schwertlilie zugehörig (lat.), mit Diminutiv; möglicherweise auch Bezug zum weiblichen Vornamen Iris Issus F. Kilikische Hafenstadt nahe Iskenderum, berühmt durch die Schlacht Alexanders des Großen gegen den Perserkönig Darius Japananus Ball der Japaner (nlat.) Jassargus Zachv. Kombination aus Jassus und argus (siehe dort)

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Jassidaeus Fieb. der dem Jassus ähnliche (siehe dort) Javesella Fenn. die kleine Javesa (hebr.: dürr), ein von Amyot verwendetes, nomenklaturisch ungültiges Mononym Jikradia Nielson akronymisch kombiniert aus Jim Kramer und der Endung der revidierten Gattung Coelidia Kelisia Fieb. die mit einer Schere ausgestattete (griech.: chelé), bezogen auf die Pygophoranhänge Kervillea Bgv. benannt nach dem französischen Entomologen und Naturforscher Henri Gadeau de Kerville (1858–1940) Kosswigianella W. Wg. nach dem Hamburger Zoologen Curt Kosswig (1903–1982) benannt Kyboasca Zachv. von Kybos (siehe dort) und Empoasca (siehe dort) Kybos Fieb. der Würfel (griech.), vermutlich bezogen auf den eckigen Vorderkörper Laburrus Rib. von labur (bask.: kurz) Lamprotettix Rib. die Glanzzirpe (griech.) Laodelphax Fenn. eventuell kombiniert aus Volk (griech.) und delphax (siehe dort), vermutlich bezogen auf die Häufigkeit Latematium Dlab. Kombination aus latus (lat.: breit) und Agalmatium (siehe dort) Latissus Dlab. der breite (lat.) Issus (siehe dort) Lausulus Rib. von lauso (bask.: neblig, trübe), bezogen auf das hyaline Flügelfenster Lebradea Rem. nach dem locus typicus Lebrade in Ostholstein benannt Ledra F. vorgriech. Name der zyprischen Stadt Lefkosia Leimonodite Kirk. die auf der Wiese geborene (griech.) Leopallia Gnezd. nach dem Beschreiber der Typusart, dem tschechischen Arzt und Zikadologen Leopold Melichar (1856–1924) benannt Lepyronia Am. et Serv. frei: die Eierschalenartige (griech.), bezogen auf die Körperform Liburnia Stål nach Liburnien, einem Küstenlandstrich im heutigen Kroatien benannt Liguropia Hpt. von Ligurien (Küstenlandschaft in Nordwest-Italien) und -opia (siehe unter Paropia) Limotettix J. Shlb. die Sumpfzirpe (lat./griech.) Lindbergina Dlab. nach dem finnischen Hemipterologen Håkan Lindberg (1898–1966) benannt Linnavuoriana Dlab. nach dem finnischen Hemipterologen Rauno Linnavuori (*1927) benannt Litemixia Asche nach dem locus typicus Lit-e-Mixe in Südwest-Frankreich benannt Lonatura Ball unklar, vermutlich eine willkürliche Neukombination Lyristes Horv. der Leierspieler (griech.)

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Macropsidius Rib. der Macropsis ähnliche (siehe dort) Macropsis Lew. das große Gesicht (griech.) Macrosteles Fieb. mit langem Stiel (griech.), bezogen auf den langen verschmolzenen Abschnitt von Radius und Media im Vorderflügel Macustus Rib. vermutlich von mako (bask.: Haken), bezogen auf die Penisform Maiestas Dist. Majestät (lat.), bezogen auf die kronenartige Zeichnung des Kopfvorderrandes Malenia Hpt. Anagramm von Lamenia Stål Megadelphax W. Wg. das große Ferkel (griech.) Megamelodes Le Q. der dem Megamelus ähnliche (siehe dort) Megamelus Fieb. von mega (griech.: groß) und melos (griech.: Glied), bezogen auf das vergrößerte Antennengrundglied Megophthalmus Curt. das große Gesicht, eventuell auch: das große Auge (griech.) Melampsalta Kol. die schwarze Sängerin (griech.) Membracis F. von membrana (griech.: Pergament), bezogen auf die Flügel Mendrausus Rib. von mendratu (bask.: verschleißen, abnutzen), bezogen auf das stärker eingeschnittene 7. Abdominalsternit der Weib- chen Metadelphax W. Wg. inmitten von Delphax (siehe dort) Metalimnus Rib. inmitten des Sumpfes (griech.) Metcalfa Caldw. nach dem amerikanischen Hemipterologen Zeno Payne Metcalf (1885–1956) benannt Metidiocerus Oss. vermutlich: dem Idiocerus nahe (griech.), siehe dort Metropis Fieb. eventuell von metron (griech.: Maß, Maßstab) und tropis (griech.: Schiffskiel), Bezug jedoch unklar Micantulina An. vom älteren Artnamen micantula gebildet (siehe dort) Mimallygus Rib. der Nachahmer von Allygus, siehe dort (griech./lat.) Mirabella Em. nach der eitlen Prinzessin aus Andersens Märchen “Die Prinzessin auf der Erbse” Mocuellus Rib. vermutlich von moko (bask.: Schnabel), bezogen auf die Penisform Mocydia Edw. eventuell: die durch Ehebruch erzeugte (griech.), Bezug jedoch unklar Mocydiopsis Rib. die wie Mocydia aussehende (griech.) Mongolojassus Zachv. der mongolische Jassus (siehe dort) Muellerianella W. Wg. nach dem ostdeutschen Zoologen und Entomologen Hans Joachim Müller (1911–2007) benannt Muirodelphax W. Wg. nach dem hawaiianischen Entomologen Frederick Arthur Godfrey Muir (1872–1931) benannt, delphax siehe oben Mycterodus Spin. von mycter (griech.: Nüster) Myndus Stål griech. Hafenstadt in Karien, heute Südwest-Türkei

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Nanocixius W. Wg. der zwergenhafte (lat.) Cixius (siehe dort) Nanosius Dlab. von nanus (lat./griech.: Zwerg ) Neoaliturus Dist. wörtlich: der neue, andersartige Schwanz Neocixius W. Wg. der neue Cixius – siehe dort Neophilaenus Hpt. der neue Philaenus (siehe dort) Nesoclutha Evans von nesos (griech.: Insel) und Balclutha (siehe dort), da aus Australien beschrieben Nothodelphax Fenn. das unechte Ferkel (griech.) Notus Fieb. vermutlich von noton (griech.: der Rücken ), bezogen auf die Wölbung des Pronotums Oliarus Stål griechische Insel, den Kykladen zugehörig Ommatidiotus Spin. der mit eigenartigen Augen (griech.) Ommatissus Fieb. wörtlich: der Augen-Issus (griech.) Oncodelphax W. Wg. mit Haken versehenes Ferkel (griech.), bezogen auf die Form der Styli Oncopsis Burm. das geschwollene Gesicht (griech.) Ophiola Edw. vermutlich von ophis (griech.: Schlange), vermutlich bezogen auf die mitunter schlängelartige Marmorierung des Pronotums Opsius Fieb. der mit großen Augen oder großem Gesicht versehene (griech.) Orientus De L. der Orientale (lat.) Orinocixius W. Wg. der Gebirgs-Cixius (griech.) – siehe dort Ossiannilssonola Christian nach dem schwedischen Hemipterologen Frej Ossiannilsson (1908–1995) benannt Pagiphora Horv. die eine Schlinge trägt (griech.), bezogen auf die erweiterte Radialzelle bzw. die umlaufende Ader Paluda De L. frei: die Sumpfbewohnerin (von lat.: palus = Sumpf) Pantallus Em. der gänzlich andere (griech.) Paracixius W. Wg. neben Cixius (siehe dort) Paradelphacodes W. Wg. der neben Delphacodes (siehe dort) Paradorydium Kirk. mit Dorydium verwandt (siehe dort) Paraliburnia J.-H. ähnlich Liburnia (siehe dort) Paralimnus Mats. neben dem Sumpf (griech.) Paramesus Fieb. wörtl.: neben der Mitte (griech.), vermutlich bezogen auf ein Kopfmerkmal Parapotes Em. der am Strom lebende (nach Emeljanov, pers. Mitt.) Paropia Germ. vermutlich von pareia (griech.: Wange) und opsis (griech.: Auge, Gesicht) Pastiroma Dlab. kombiniert aus pa- (tschech.: pseudo-) und Stiroma (siehe dort) Pediopsis Burm. das große, flache Gesicht (griech.) Peltonotellus Put. der kleine Peltonotus (siehe dort) Peltonotus M. et R. von pelte (griech.: Schild) und notos (griech.: Rücken)

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Penestragania fast Stragania (griech.: strangaliaô = einschnüren), bezogen Beam. et Laws. auf die Skulpturierung des Pronotums Pentastira Kbm. die Fünfkielige (griech.) Pentastiridius Kbm. ähnlich der Pentastira (siehe oben) Penthimia Germ. die Trauernde (griech.) Perotettix Rib. die Stiefelzirpe (lat./griech.), bezogen auf die Penisform Peuceptyelus J. Shlb. von peuke (griech.: Nadelbaum) und ptyelon (griech.: Speichel) Phantia Fieb. die Sichtbare (griech.) Philaenus Stål unklar: von philenor (griech.: verliebt); oder: einer von zwei Brüdern aus Karthago, die sich aus Vaterlandsliebe lebendig begraben ließen; oder: der die (Lob-)Rede liebende Philaia Dlab. frei: die liebe, nette (griech.) Phlepsius Fieb. von phleps (griech.: Ader), bezogen auf die stark geaderten Flügel Phlogotettix Rib. die feurige Zirpe (griech.) Pholetaera Zett. von pholeter (griech.: der in einer Höhle lauert) Pinumius Rib. vermutlich von Pinus (lat./bask.: Kiefer), Bezug unklar Pithyotettix Rib. die Fichtenzirpe (griech.) Placotettix Rib. Plattenzirpe (griech.), vermutlich bezogen auf die Form des Konnektivs Planaphrodes Ham. die abgeplattete Aphrodes (siehe dort) Platymetopius Burm. mit abgeplatteter Stirn (griech.) Pleargus Em. kombiniert aus pleos (griech.: klein) und Jassargus (siehe dort) Polania Em. vermutlich nach dem Stamm der Poljanen, in deren Gebiet in der heutigen Ukraine der locus typicus liegt Populicerus Dlab. frei: der mit Hörnern versehene auf der Pappel (lat./griech.) Praganus Dlab. frei: der Prager (nlat.) Proceps M. et R. der vorstehende Kopf (lat.) Prokelisia Osb. aus Kelisia (siehe dort) herausragend Psammotettix Hpt. die Sandzirpe (griech.) Pseudodelphacodes W.Wg. der wie Delphacodes erscheinende (siehe dort) Le P. et Serv. von ptyelon (griech.: Speichel) Recilia Edw. vermutlich von cilia (lat.: Wimpern, Lider), vermutlich bezogen auf die nahe der Komplexaugen liegenden Ozellen Reptalus Em. frei: der kriechende Geflügelte (lat.) Rhoananus Dlab. unklar, eventuell aus Sorhoanus umgebildet und verändert (siehe dort) Rhopalopyx Rib. frei: der Keulensteißige (griech.) Rhytidodus Fieb. frei: der mit Runzeln versehene (griech.) Rhytistylus Fieb. der runzelige Fortsatz (griech.), vermutlich bezogen auf den Kopfvorderrand

284 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen

Ribautiana Zachv. nach dem französischen Hemipterologen und Myriapodo- logen Henri Ribaut (1872–1967) benannt Ribautodelphax W. Wg. nach Henri Ribaut benannt (siehe oben); delphax siehe dort Rosenus Oman unklar, vermutlich eine willkürliche Neukombination Sagatus Rib. der Weidenbewohner (bask.: sagatz = Weide) Sahlbergotettix Zachv. nach dem finnischen Zikadologen und Koleopterologen Johan Reinhold Sahlberg (1845–1920) benannt; tettix = Zirpe (griech.) Sardius Rib. frei: der Gegabelte (bask. sarde = Forke), bezogen auf die Form des Konnektivs Sayetus Rib. möglicherweise von sai (bask.: Aasgeier), bezogen auf die Penisform Scaphoideus Uhl. der Kahnartige (griech.) Sciocixius W. Wg. der dunkle Cixius (griech.) – siehe dort Scleroracus Van Duz. frei: der rauhe Lumpen trägt (griech.) Scottianella An. nach dem englischen Entomologen John Scott (1823–1888) benannt Selenocephalus Germ. der Mondkopf (griech.) Setapius Dlab. von seta (lat.: Borste) und pius (lat.: freundlich, gerecht) Sonronius Dorst benannt nach dem Sohn des Beschreibers, Ronald A. Dorst (engl.: „son Ron“) Sorhoanus Rib. der Graslandbewohner (bask.: sorho = Wiese, Weide) Sotanus Rib. der Höhlenbewohner (span. oder bask.) Speudotettix Rib. vermutlich: speudo = Anagramm zu pseudo; tettix = Zirpe (griech.) Stenidiocerus Oss. der schlanke mit besonderen Hörnern (griech.) Stenocranus Fieb. der Schmalschädel (griech.) Stenokelisia Rib. die schmale (lat.) Kelisia (siehe dort) Stictocephala Stål der punktierte Kopf (griech.) Stictocoris Thoms. die punktierte Wanze (griech.) Stiroma Fieb. die mit Kiel versehene (griech.) Stiromella W. Wg. eine kleine Stiroma (siehe dort) Stiromoides Vilb. der Stiroma ähnlich (griech./lat.) Streptanus Rib. der verdrehte Ring (lat.), bezogen auf die Penisform Streptopyx Lnv. der mit verdrehtem Steiß (griech.) Stroggylocephalus Fl. der Rundkopf (griech.) Struebingianella W. Wg. nach der Berliner Entomologin und Bioakustikforscherin Hildegard Strübing (1922–2013) benannt Hpt. frei: durch eine Braue verbundene Augen (griech.) Tamaricella Zachv. von Tamarix (lat.: Tamariske) Tetartostylus W. Wg. wörtlich: der Viertel-Stylus (lat.), bezogen jedoch auf die zwei zusätzlichen Pseudostyli Tettigetta Kol. die kleine Zikade (griech./franz.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 285 H. NICKEL

Tettigometra Latr. von Aristoteles gebrauchter Terminus für die Larven der Singzikaden, wörtlich: Mutterleib der Zikade (griech.) Thamnotettix Zett. die Strauchzikade (griech.) Tibicina Kol. die Flötenspielerin (lat.) Toya Dist. vermutlich von Wasser (sanskr.), Bezug jedoch unklar Tremulicerus Dlab. frei: der mit Hörnern versehene auf der Zitterpappel (Populus tremula) Trepalus Em. Anagramm aus Reptalus (siehe dort) Trigonocranus Fieb. der dreieckige Schädel (griech.) Tropidocephala Stål die Schiffskielköpfige (griech.) Trypetimorpha A. Costa die Bohrerförmige (griech.) Tshurtshurnella Kusn. nach dem Homopterennamen tshurtshurna (tatar.) Turrutus Rib. frei: der mit einem Turm versehene (lat. turris = Turm), bezogen auf die Penisform Typhlocyba Germ. frei: die Blindköpfige (griech.), bezogen auf die fehlenden Ozellen Tyrphodelphax Vilb. kombiniert aus tyrphe (griech.: Torf) und delphax (siehe dort) Ulopa Fall. die Narbengesichtige (griech.) Unkanodes Fenn. Ersatzname für den präokkupierten Namen Unkana Matsumura (japan.: unka = fulgoromorphe Zikade) Utecha Em. Trost, Freude (russ.) Verdanus Oman vermutlich von verde (span.: grün) Viridicerus Dlab. der grüne mit Hörnern (lat./griech.) Wagneriala An. nach dem Hamburger Hemipterologen Wilhelm Wagner (1895–1977) benannt Wagneripteryx Dlab. nach Wilhelm Wagner (siehe oben), pteryx = frei: der geflügelte (griech.) Weidnerianella W. Wg. nach dem deutschen Entomologen Herbert Weidner (1911–2009) Xanthodelphax W. Wg. das gelbe Ferkel (griech.) Ziczacella An. nach der Zickzack-Zeichnung auf den Flügeln (nlat.) Zonocyba Vilb. kombiniert aus zona (lat.: Gürtel) und Typhlocyba (siehe dort) Zygina Fieb. vermutlich: ähnlich einem Joch (griech.), vermutlich bezogen auf die Form der Randzellen der Hinterflügel Zyginella P. Löw die kleine Zygina (siehe dort) Zyginidia Hpt. kombiniert aus Zygina und Idia (siehe dort)

Liste der Arten und Unterarten abbreviatus Leth. der verkürzte (lat.) abdominalis F. das Abdomen betreffend (lat.), bezogen auf die auffällige Schwarzfärbung

286 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen abietinus Fall. der Tanne zugehörig (lat.) abrotani Em. der Eberraute (Artemisia abrotanum) zugehörig (lat.) abstrusus Lnv. der verborgene (lat.) aceris Logv. dem Ahorn zugehörig (lat.) achilleae Mity. der Schafgarbe zugehörig (lat.) acicularis M. et R. der nadelspitzige (lat.) acuminatus F. die zugespitzte (lat.) acuta P. Löw die spitze (lat.) adela Fl. die unbekannte, versteckte (griech.) adriaticus W. Wg. der adriatische (lat.) adspersa H.-S. die gesprenkelte (lat.) adumbrata C. Shlb. die schattenhafte, undeutliche (lat.) adustus H.-S. der angesengte (lat.) aemulans Kbm. der nachahmende (lat.) aestuarina Edw. die zur Flussmündung gehörige (lat.) affinis Fieb. die verschwägerte (lat.) agrestis Fall. auf dem Feld lebend (lat.) akdzhusani Zachv. dem weißen Wermut zugehörig (kasach.: ak-dzhusan) albae W. Wg. der Silber-Weide (Salix alba) zugehörig (lat.) albicans Kbm. der weiß schimmernde (lat.) albifrons L. weiße Stirn (lat.) albiger Germ. der Weißträger (lat.) albingensis W. Wg. von der Region Unterelbe stammend (lat.: Albingia) albipennis F. mit weißen Flügeln (lat.) albocarinata Stål die weißgekielte (lat.) alboguttatus Leth. der weiß betropfte (lat.) albomarginatus Curt. der weißrandige (lat.) alboniger Leth. der weißschwarze (lat.) albosignata Dhlb. die weiß bezeichnete (lat.) albostriata Fieb. die weißgestreifte (lat.) albostriella Fall. vermutlich: die weiß gestrichelte (lat.) alces Rib. der Elch (lat.), bezogen auf die Aedeagusform alemannicus W. Wg. der alemannische (lat.) alienus Dhlb. der fremde (lat.) allionii Turt. nach dem Turiner Botaniker Carlo Allioni (1728–1804) benannt allobrogicus Rib. der allobrogische (lat.), bezogen auf ein Gebiet in der heutigen Provence alneti Dhlb. dem Erlenwald zugehörig (lat.) alni F. der Erle zugehörig (lat.) alnicola Edw. der Erlenbewohner (lat.) alnisuga Arz. der Erlensauger (lat.) alpestris W. Wg. das Hochgebirge bewohnend (nlat.) alpicola W. Wg. der Alpenbewohner (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 287 H. NICKEL alpinus Zett. dem Gebirge zugehörig (lat.) alsiosa Rib. die frostempfindliche (lat.) alsius Rib. vermutlich: der kühle (lat.) altaica Vilb. die aus dem Altai stammende (nlat.) alticola Rib. der Höhenbewohner (lat.) ampliata W. Wg. die vergrößerte (lat.), vermutlich bezogen auf die im Vergleich mit E. frustrator (Edw.) verlängerten Aedeagusanhänge anatolica Zachv. die anatolische (lat.) anceps Germ. der doppelköpfige, zweiseitige (lat.), Bezug unklar andalusiaca Ferr. die andalusische (lat.) anderi Oss. nach dem schwedischen Entomologen Kjell Ander (1902–1992) andropogonis Horv. dem Bartgras (Andropogon) zugehörig (nlat.) angulatus Then der eckige (lat.) angulosa Rib. die eckige (lat.), bezogen auf die Ausbuchtung auf den Styli angusta Leth. die schmale (lat.) annulata Br. die beringte (lat.) antennalis Hpt. der mit besonderen Antennen versehene (lat.) antoniae Mel. nach Leopold Melichars Ehefrau Antonia benannt apicalis Fl. frei: mit einer auffälligen Spitze versehen (lat.), bezogen auf die Flügel apiculatus Fieb. der fein zugespitzte (lat.) appendiculata W. Wg. die mit Anhang versehene (lat.) apterus F. der ungeflügelte (griech.) arctous Em. der nördliche (lat.) arcuata Fieb. die bogenförmig gekrümmte (lat.) areatus Stål der getäfelte (lat.), Bezug unklar arenarius Rem. zum Sand gehörig (lat.) argentarius Metc. der silberne (lat.) argus Marsh. hundertäugiger Bewacher der Io, einer Geliebten des Zeus arhenana Dlab. wörtlich: die unrheinische, bezogen auf die Unterschiede zur ähnlichen rhenana W.Wg. (s. dort) aridella J. Shlb. die etwas dürre (lat.) artemisiae Kbm. dem Wermut zugehörig (lat.) asemus Fieb. der unkenntliche (griech.) asper Rib. der rauhe (lat.) assimilis Fall. der ziemlich ähnliche (lat.) aterrimus J. Shlb. der sehr schwarze (lat.) atomarius F. der ungeteilte (lat.) atra Hag. die schwarze (lat.) atrata Fieb. die schwarz gekleidete (lat.) atricapillus Boh. der schwarzhaarige (lat.)

288 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen atropidicola Em. Bewohner des Salzschwadens (Atropis = Puccinellia; nlat.) atropunctata Goeze die schwarz gepunktete (lat.) attenuata Germ. die verschmälerte (lat.) aubei Perr. nach dem französischen Entomologen Charles Aubé (1802–1869) benannt audrasi Rib. nach dem Lyoner Koleopterologen Georges Audras (1881–1970) benannt aurantiacus Forel der organgerote (lat.) aurantipes Edw. die goldfüßige (lat.) aurata L. die vergoldete (lat.) aureola Fall. die goldene (lat.) aurita L. die geöhrte (lat.) aurovittata Dgl. die golden gestreifte (lat.) aurulentus Kbm. der goldfarbene (lat.) austriaca W. Wg. die österreichische (nlat.) avellanae Edw. der (Hasel-)Nuss zughörig, benannt nach der kampanischen Stadt Abella (lat.) aviger Em. der Vogelträger (lat.), benannt nach der Scheitelzeichnung axillaris J. Shlb. die durch ihre Schulter ausgezeichnete (lat.) baeri Kouch. nach dem deutsch-baltischen Naturforscher Karl Ernst von Baer (1792–1876) benannt baldensis Rem. et Sch. vom norditalienischen Monte Baldo stammend (nlat.) baranii Sign. nach dem Sammler des Holotypus, Gabriel de Baran (1830–1864) benannt barbata Rib. die bärtige (lat.) basilinea Germ. Strich an der Basis (lat.) beckeri Fieb. nach dem sareptischen Biologen Alexander Becker (1818–1901) benannt beieri W. Wg. nach dem österreichischen Arachnologen und Entomologen Max Beier (1903–1979) benannt bellevoyei Put. nach dem Lothringer Entomologen und Künstler Adolphe Nicolas Bellevoye (1830–1908) benannt bensoni China nach dem britischen Entomologen Robert Bernard Benson (1904–1967) benannt beresi Orosz nach dem rumänischen Entomologen József Béres bergmani Tull. nach dem schwedischen Veterinär, Rentierforscher und Sammler des Holotypus Arvid Mathias Bergman (1872–1923) benannt bicarinata H.-S. die mit zwei Kielen versehene (lat.) bicincta Schrk. die mit zwei Gürteln versehene (lat.), bezogen auf die hellen Querbänder bicolor Logv. die zweifarbige (lat.) bicorniger Then der zwei Hörner tragende (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 289 H. NICKEL bicuspidatus J. Shlb. der zweifach bestachelte (lat.) bielawskii Nast nach dem polnischen Koleopterologen Ryszard Bielawski (*1930) benannt bifasciata L. die zweifach gebänderte (lat.) bilobum Fieb. das zweilappige (lat./griech.) binotatus J. Shlb. der zweifach gekennzeichnete (lat.) bipunctata Osh. die zweifleckige (griech./lat.) bisignata M. et R. die zweifach gezeichnete (lat.) bisonia Kopp et Yonke wörtlich: der auerochsenartige (lat.), die Autoren bezogen sich aber sicherlich auf den amerkanischen Bison bispinatus Then der zweifach bedornte (lat.) bisubulatus Then der zweispitzige (lat.) blandula Rossi die kleine reizende (lat.) bobbicola Rem. et Sch. Bewohner des norditalienischen Monte Bobbio bohemani Zett. nach dem schwedischen Zoologen Carl Henrik Boheman (1796–1868) benannt bohemica Dlab. die böhmische (lat.) boica W. Wg. die bayerische (lat.) boldi Scott nach dem britischen Entomologen Thomas John Bold (1816–1874) benannt bonellii Latr. nach dem italienischen Zoologen Franco Andrea Bonelli (1784–1830) benannt borealis J. Shlb. der nördliche (lat.) borussicus W. Wg. der preußische (nlat.) bosnica Horv. die bosnische (nlat.) bottnica Huldén der bottnische (lat.) brabantica W. Wg. die brabantische (nlat.) brachyanus Orosz der kurzsteißige (griech./lat.) brachycranus Scott der kurzschädelige (griech.) brachynota Fieb. die kurzrückige (griech.) brachyptera Boh. die kurzflügelige (lat.) brevipennis Fieb. mit kurzen Flügeln (lat.) brevis H.-S. der kurze (lat.) bromi Em. der Trespe (Bromus) zugehörig brucki Fieb. nach dem Krefelder Koleopterologen und Kommerzienrat Emil vom Bruck (1807–1884) benannt brullei Fieb. nach dem französischen Entomologen Gaspard Auguste Brullé (1809–1873) brunnea Mel. die braune (lat.) butleri Edw. nach dem englischen Entomologen, Arachnologen und Ornithologen Edward Albert Butler (1844–1925) benannt calamagrostidis Rem. dem Reitgras (Calamagrostis) zugehörig (nlat.) calamagrostis Oss. wie oben

290 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen calcarata Oss. die gespornte (lat.) caledonicus China der schottische (lat.) callosa Then die dickhäutige (lat.) calyculus Cer. kleiner Blütenkelch (lat.), bezogen auf die Penisform cambricus China der walisische (lat.) campestris Fall. der Ebene zugehörig (lat.) candidula Kbm. die schön weiße (lat.) cantilatrix Sueur et Puiss. die Sängerin (lat.) capicola Stål der Kap-Bewohner (lat.) capnodes Scott der rauchige (griech.) carinata Stål die gekielte (lat.) carneae W. Wg. der Schneeheide (Erica carnea) zugehörig carniolicus W. Wg. der Krainer (lat.) carpathica Mel. die karpatische (lat.) carpini J. Shlb. der Hainbuche zugehörig (lat.) carri Edw. nach dem britischen Botaniker und Entomologen John Wesley Carr (1862–1939) benannt caudatus Fl. der mit Schwanz versehene (lat.) cephalotes H.-S. durch den Kopf ausgezeichnet (griech.) cerasi Em. der Kirsche zugehörig (lat.) cerdaniensis Puiss. et Boul. nach dem locus typicus Cerdagne in den französischen Pyrenäen cerea Germ. vermutlich: die wachsfarbene (lat.) cerricola Laut. der Bewohner der Zerr-Eiche (Quercus cerris; nlat.) chinai Oss. nach dem britischen Hemipterologen William Edward China (1895–1979) chloris An. von chlorós (griech.: grün) chloroticus Mel. der mit Grün versehene (lat.) chobauti Rib. nach dem französischen Koleopterologen Alfred Chobaut (1860–1926) benannt ciliata Osb. die bewimperte (lat.) cinerascens Em. der aschgrau werdende (lat.) cisalpina Dwor. die diesseits der Alpen lebende (lat.) citrinella Zett. die zitronengelbe (lat.) clavata Dlab. die mit Keulen versehene (lat.) clavicornis F. mit keulenförmigen Hörnern (lat.) clypealis J. Shlb. die mit einem besonderen Schild versehene (lat.) clypeata Horv. wie oben cognatus Fieb. der Blutsverwandte (lat.) coleoptrata L. die käferartige (nlat.) collina Boh. die auf dem Hügel lebende (lat.) comitans Em. der begleitende (lat.) communis Ferr. der allgemeine (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 291 H. NICKEL commutatus Fieb. der veränderte (lat.) complicatus Nast der verworrene (lat.) concinna Germ. die zierliche, feine (lat.) concolor Fieb. die einfarbige (lat.) concors Horv. die übereinstimmende (lat.) confinis Dhlb. der verwandte (lat.) confusus Fl. der undeutliche (lat.) conica Say die kegelförmige (lat.) consobrina Curt. die Cousine (lat.) conspurcatus Perr. der besudelte (lat.) contraria Rib. die Gegnerin (lat.) convenarum Rib. wörtlich: der zusammen kommenden (lat.), Bezug unklar coriaceus Fall. der lederne (lat.) cornicula Marsh. kleine Hörner (lat.) cornutus L. der gehörnte (lat.) coronifera Marsh. der Kranzträger (lat.) corticea Germ. die zur Rinde gehörige (lat.) corvina Horv. die rabenartige (lat.), bezogen auf die Färbung coryli Le Q. der Hasel zugehörig (lat.) costalis Fall. mit besonderem (Flügel-) Rand (lat.), bezogen auf die Hell- Dunkel-Zeichnung costata Curt. die gerippte (lat.) crassicornis Panz. mit dickem Horn versehen (lat.) crassipes J. Shlb. der dicke Fuß (lat.) crataegi Dgl. dem Weißdorn zugehörig (lat.) cristatus Rib. der mit Kamm versehene (lat.) crocea H.-S. die safranfarbige (lat.) cruciata Rib. die mit einem Kreuz versehene (lat.), bezogen auf den kreuzförmig verzweigten Aedeagusschaft cruenta H.-S. die blutrote (lat.) cruentatus Panz. der mit Blut bespritzte (lat.) cuniceps Mel. der keilköpfige (lat.) cunicularius L. der Minierer (lat.); Bezug unklar, die epigäische Lebensweise der Larven war Linné vermutlich nicht bekannt cupreus Kbm. der kupferfarbene (lat.) curtisii Fl. nach dem englischen Entomologen und Naturmaler John Curtis (1791–1862) benannt curvatus Dlab. der gekrümmte (lat.) cuspidata F. die gestachelte (lat.) cyane Boh. die schwarzblaue (griech.) cyclops Mats. der Zyklop (lat.), vermutlich bezogen auf den unpaaren Scheitelfleck cytisi Zachv. der dem Geißklee zugehörige (lat.)

292 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen dahlbomi Zett. nach dem schwedischen Entomologen Anders Gustav Dahlbom (1808–1859) benannt dalei Scott nach dem britischen Entomologen Charles William Dale (1791–1872) benannt dalmatina W. Wg. die dalmatische (lat.) dealbata Cer. die getünchte (lat.) debilis Dgl. die gebrechliche (lat.) decedens Paoli die weggehende (lat.) decemnotata R. die zehnfach gekennzeichnete (lat.) decempunctata Fall. die mit zehn Punkten versehene (lat.) decimusquartus H.-S. der vierzehnte (lat.), bezogen auf das diagnostische Zeich- nungselement im Flügel decipiens Paoli die täuschende (lat.) decumana Kontk. die ansehnliche (lat.) dehneli Nast nach dem polnischen Theriologen August Dehnel (1903–1962) benannt dentatus D’Urso der mit einem Zahn versehene (lat.) denticauda Boh. der Zahnschwanz (lat.) depressa Fieb. die niedergedrückte (lat.) desertorum Fieb. der öden Gegenden (lat.) detectus Rib. der entdeckte (lat.) digitatus Rib. der gefingerte (lat.) dilatatus Fourcr. der verbreiterte (lat.) dilutior Kbm. der schwächer gefärbte (lat.) dimidiatus Rib. der halbierte (lat.) diminuta Rib. die verkleinerte (lat.) dimissa Hagen die weggeschickte (lat.) discolor Boh. die andersfarbige (lat.) discrepans Fieb. die abweichende (lat.) dispar Fieb. der ungleiche (lat.) dissimilis Fall. der unähnliche (lat.) distincta Fl. die andersartige (lat.) distinguendus Kbm. der zu unterscheidende (lat.) divergens Kbm. die auseinandergehende (lat.), bezogen auf die Stirnkiele diversa Edw. die auseinander gekehrte (lat.), bezogen auf die Aedeagusanhänge dlabolai W. Wg. nach dem böhmischen Zikadologen Jiøí Dlabola (*1922) benannt domino Reut. der Herr (lat.) dorsatum Ahr. das mit besonderem Rücken versehene (lat.) dorsofenestratus Dlab. der mit Fenstern auf dem Rücken versehene (lat.) douglasi Scott nach dem englischen Mikrolepidopterologen und Hemipterologen John William Douglas (1814–1905)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 293 H. NICKEL dubia Kbm. die zweifelhafte (lat.) duffieldi Le Q. nach dem englischen Entomologen C.A.W. Duffield (1887–1974) benannt dumosa Rib. die verzweigte, heckenartige (lat.), auf die Penisform bezogen duodecimguttatus Cer. der zwölffach betropfte (lat.) elaeagni Em. der Ölweide zugehörig (lat.) elegans Fl. der feine, geschmackvolle (lat.) elegantula Boh. die niedliche (lat.) elongata W. Wg. die verlängerte (lat.) emmeae Fieb. nach dem schweizerischen Fluss Emme benannt empetri Oss. zur Krähenbeere gehörend (nlat.) erecta Rib. die aufrechte (lat.), bezogen auf die Form der Styli ernesti Günth. nach dem Schweizer Phytopathologen Ernst Günthart (1917–1990) benannt erraticus Lnv. der auf Abwegen gehende (lat.) erythrocephala Ferr. die rotköpfige (griech.) estonica Vilb. die estnische (nlat.) etnicola Asche der Ätna-Bewohner (lat.) eugeniae Kusn. nach der Gattin des Erstbeschreibers, Eugenie Kusnetsov, benannt europaea L. die europäische (lat.) evanescens Oss. die verschwindende (lat.) excisus Mats. der eingeschnittene (lat.), Bezug unklar exclamationis Thnbg. des Ausrufes (lat.), bezogen auf die weiße Flügelzeichnung, die einem Ausrufezeichen ähnelt exemtus Mel. der herausgenommene (lat.) exigua Boh. die winzige (lat.) exilis Horv. die winzige, dürftige (lat.) eximius Kbm. der besondere (lat.) exornatus Rib. der ausgestattete (lat.) extrusa Scott die verstoßene (lat.) fairmairei Perr. nach dem französischen Koleopterologen Léon Fairmaire (1820–1906) benannt falleni Metc. nach dem schwedischen Entomologen Carl Fredrik Fallén (1764–1830) benannt fasciatus Fieb. der gebänderte (lat.) fasciolatus Mel. der mit feinem Band versehene (lat.) fenestratus H.-S. der mit Fenstern versehene (lat.), bezogen auf die Flügelfärbung fennahi Young nach dem britischen Zikadologen Ronald Gordon Fennah (1910–1987) benannt ferrarii Put. nach dem italienischen Hemipterologen und Arzt Pietro Mansueto Ferrari (1823–1893) benannt

294 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen ficaria Horv. zur Feige (lat.: ficus) gehörig fieberi Edw. nach dem böhmischen Hemipterologen Franz Xaver Fieber (1807–1872) benannt filicum Newm. den Farnen zugehörig (lat.) flammigera Fourcr. die Flammenträgerin (lat.) flava P. Löw die gelbe (lat.) flaveola Boh. die gelbliche (lat.) flavescens F. gelb werdend (lat.) flaviceps Fieb. der gelbköpfige (lat.) flavicollis L. die gelbhalsige (lat.) flavida Vilb. die gelbliche (lat.) flavipennis Zett. der gelbflügelige (lat.) flavipes Sign. der Gelbfuß (lat.) flavobrunnea Dlab. die gelbbraune (lat.) flavostriatus Don. der gelbgestreifte (lat.) flavovarius H.-S. der gelb-bunte (lat.) flavovirens Gill. et Bak. die gelbgrün werdende (lat.) flexuosa Rib. die gewundene (lat.), nach der Form der Aedeagusanhänge flori Fieb. nach dem livländischen Hemipterologen Gustav August Adam von Flor (1829–1883) benannt florii Stål wie oben, aber vom latinisierten Namen Florius gebildet florida Rib. wörtlich: die blühende (lat.), aber auch Bezug auf G.A. Flor (siehe oben) forcipata Boh. die mit einer Zange versehene (lat.) forficula Rib. die kleine Schere (lat.) formosus Boh. der schöne (lat.) fortior W. Wg. die stärkere (lat.) foveolatus Fieb. der mit einer Grube versehene (lat.) fragilicola Bewohner der Bruch-Weide (Salix fragilis) Hlz., Nick. et Rem. franzi W. Wg. nach dem österreichischen Entomologen und Hochgebirgs- ökologen Herbert Franz (1908–2002) benannt fraudulenta Horv. die trügerische (lat.) frauenfeldi Leth. nach dem österreichischen Entomologen und Malakologen Georg Ritter von Frauenfeld (1807–1873) benannt freyi Fieb. nach dem Schweizer Hemipterologen Emil Frey-Gessner (1826–1917) benannt frisia W. Wg. die friesische (lat.) frontalis H.-S. die durch ihre Stirn gekennzeichnete (lat.) frustrator Edw. der Täuscher (lat.) fruticola Fall. der Strauchbewohner (lat.) fuchsii Kbm. nach dem Schmetterlingssammler Georg Karl August Ferdinand Fuchs (1839-1904), Pfarrer im Nassauischen Bornich, benannt

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 295 H. NICKEL fulgidus F. der glänzende (lat.) fumipennis Fieb. mit rauchfarbenem Flügel (lat.) furcatus Ferr. der gegabelte (lat.) fusca Fieb. die schwärzliche (lat.) fuscinervis Boh. die dunkelnervige (lat.) fuscofasciatus Goeze der dunkel gebänderte (lat.) fuscovenosus Ferr. der dunkel geaderte (lat.) fuscovittatus Stål der dunkel gestreifte (lat.) fuscula Zett. die bräunliche (lat.) gallica W. Wg. die gallische (lat.) genei Spin. nach dem Sammler des Holotypus, “M. Géné”, benannt; vermutlich handelt es sich um den Turiner Zoologen Guiseppe Gené (1800–1847) genistae F. dem Ginster zugehörig (lat.) geometrica Schrk. die geometrische (lat.) germari Zett. nach dem Hallenser Entomologen und Mineralogen Ernst Friedrich Germar (1786–1853) benannt gilveolus Kbm. der honiggelbe (lat.) glandacea Fieb. die einer Eichel ähnliche (lat.) glaucescens Fieb. blaugrün werdend (lat.) graminea F. die grasartige (lat.), vermutlich bezogen auf die Farbe grandii Serv. nach dem italienischen Entomologen (v.a. Hymenopterologen) Guido Grandi (1886–1970) gratiosa Boh. die angenehme (lat.) gravesteini W. Wg. nach dem niederländischen Hemipterologen Willem Hendrik Gravestein (1906–1989) benannt grisea Walk. die graue (lat.) griseola Fieb. die gräuliche (lat.) griseombra Rem. wörtlich: die grau beregnete (lat./griech.) grisescens Zett. grau werdend (lat.) grylloides F. der heuschreckenartige (lat.) guttatus Fieb. der gesprenkelte (lat.) gutticollis Horv. mit betropftem (= geflecktem) Hals (lat.) guttula Germ. das Tröpfchen (lat.) guttulifera Kbm. die Tröpfchen tragende (lat.) guttulinervis Kbm. mit betropften Flügeladern (lat.) gvosdevi Mity. vermutlich nach dem kasachischen Zoologen Evgeny Gvozdev (1918–2012) gyllenhalii Fall. nach dem schwedischen Koleopterologen Leonard Gyllenhaal (1752–1840) benannt haagi Kbm. nach dem Frankfurter Landwirt und Koleopterologen Georg Johann Haag von Rutenberg (1830 – 1888) benannt haematoceps M. et R. der blutrote Kopf (griech.)

296 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen haematodes Scop. blutig (griech.) hagemini Rem. et J. Verschmelzung aus haupti (siehe dort) und gemini (lat.: die Zwillinge); außerdem Bezug zum kleinen Genitalsegment (“ge-” und “-mini”) haglundi J. Shlb. nach dem schwedischen Hemipterologen Carl Johan Emil Haglund (1837–1901) benannt halpina Rem. et J. Verschmelzung aus haupti und alpina (siehe dort) hamata Boh. die mit Haken versehene (lat.) handlirschi Mats. nach dem Wiener Entomologen und Paläontologen Anton Peter Josef Handlirsch (1865–1935) benannt hardei Dlab. nach dem deutschen Koleopterologen Karl Wilhelm Harde (1922–1982) harpago Rib. der Enterhaken (lat.) harrisoni W. Wg. nach dem britischen Hemipterologen George Heslop- Harrison (1911–1964) benannt hartigi W. Wg. nach dem Südtiroler Lepidopterologen Friedrich von Hartig (1900–1980) benannt haupti W. Wg. nach dem Hallenser Homopterologen Hermann Haupt (1873–1959) benannt helva Arz. die isabellfarbene (lat.) helvolus Kbm. der gelbliche (lat.) hemiptera O. Costa die halb geflügelte (griech.) henribauti Dlab. nach dem französischen Hemipterologen und Myriapodologen Henri Ribaut (1872–1967) benannt henschii Horv. nach dem slowakischen Arzt und Entomologen Andrija Hensch (1857–1930) benannt heptapotamica Kusn. die aus dem Siebenstromland (größtenteils Kasachstan) stammende (griech.) herrichii Kbm. nach dem Regensburger Entomologen Gottlieb August Wilhelm Herrich-Schäffer (1799–1874) benannt heterophyla Horv. von anderem Stamm (griech.) heydenii Kbm. nach dem Frankfurter Entomologen und Politiker Carl Heinrich Georg von Heyden (1793–1866) benannt hieroglyphicus Kusn. der mit einem Buchstaben markierte (lat.) hippocastani Edw. der Rosskastanie (Aesculus hippocastanum) zugehörig (nlat.) hispidus Logv. der borstige (lat.) histrionicus F. der schauspielernde (lat.) hofferi Dlab. nach dem böhmischen Hymenopterologen Augustin Hoffer (1910–1981) benannt homophyla Fl. aus gemeinsamem Stamm (griech.) horvathi Then nach dem ungarischen Hemipterologen Géza Horváth (1847–1937) benannt

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 297 H. NICKEL humilis Horv. der niedrige (lat.) hungarica Rib. die ungarische (lat.) hyalinus Osb. der gläserne (lat.) hyperici H.-S. dem Johanniskraut zugehörig (lat.) hypermaculata die übermäßig gefleckte (griech./lat.), auch akronymischer Rem. et Hlz. Bezug zur Wirtspflanze Hypericum maculatum hypochlorus Fieb. der weniger grüne (griech.) identica Tish. die gleiche (nlat.) ignoscus Mel. der unbekannte (lat.) illyricus Kbm. der illyrische (lat.) imitans Rib. die ähnliche (lat.) imitantoides Biem. die der imitans (siehe dort) ähnliche immaculatifrons Kbm. die ungefleckte Stirn (lat.) immaculatus F. der ungefleckte (lat.) impictifrons Boh. die ungezeichnete Stirn (lat.) impressifrons Kbm. mit eingedrückter Stirn impressopunctata Duf. mit eingedrückten Punkten versehen (lat.) impudica Horv. die unzüchtige (lat.), Bezug unklar impura Boh. die unreine (lat.) incisus Kbm. der eingeschnittene (lat.), bezogen auf das auf der Dorsal- seite eingeschnittene Genitalsegment der ♂ incisa Then die eingeschnittene (lat.), bezogen auf den Hinterrand des 7. Abdominalsternites der ♀ inconspicuus Stål der unauffällige (lat.) inconstans Rib. die veränderliche (lat.) inermis W. Wg. die unbewaffnete (lat.), bezogen auf die fehlenden Analrohranhänge inexpectatus Rem. der unerwartete (lat.) infumatus Hpt. der eingeräucherte (lat.) infuscata J. Shlb. die angebräunte (lat.) inquinata Rib. die beschmutzte (lat.) intercedens Lnv. die dazwischen tretende (lat.) intermedia Rem. die dazwischen befindliche (lat.) interruptus L. der unterbrochene (lat.), bezogen auf den schwarzen Flügelstreifen interstinctus Fieb. der übersäte (lat.) interstitialis Germ. der dem Zwischenraum zugehörige (lat.) intractabilis Kontk. der unbehandelbare (lat.) intricatus H.-S. der verwirrte (lat.) irregulata Hpt. die nicht mit einer Leiste versehene (lat.), bezogen auf den reduzierten Stirnkiel irroratus Lew. der mit Tau benetzte (lat.) ishidae Mats. nach dem japanischen Sammler M. Ishida

298 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen italicus W. Wg. der italienische (lat.) italosignus Dros. et Rem. frei: der gezeichnete Italiener (lat.) ivanhofi Leth. nach dem Sammler M. Ivanhof ivanovi Kusn. nach dem ukrainischen Entomologen S.P. Ivanov jassargiforma Dlab. die Jassargus-artige (siehe dort) josifovi Dlab. nach dem bulgarischen Heteropterologen Michail Josifov (1927–2008) jucunda H.-S. die angenehme, die erfreuliche (lat.) juniperi Leth. dem Wacholder zugehörig (lat.) kemneri Oss. nach dem schwedischen Entomologen Nils Alarik Kemner (1887–1948), einem akademischen Lehrer von F. Ossiannilsson kirschbaumi W. Wg. nach dem Wiesbadener Hemipterologen Carl Ludwig Kirschbaum (1812–1880) benannt kobachidzei Dlab. nach dem georgischen Entomologen und Phytopathologen David Nestorowiz Kobachidze (1911–1970) benannt koeleriae Zachv. dem Schillergras (Koeleria) zugehörig (nlat.) kofleri Dlab. nach dem Osttiroler Zoologen Alois Kofler (* 1932) benannt kolenatyi Fieb. nach dem tschechisch-deutschen Naturforscher Friedrich Anton Rudolph Kolenati (1812–1864) benannt kolosvarensis Mats. der aus Kolozsvár (rumänisch: Cluj-Napoca, deutsch: Klausenburg) stammende kontkaneni Oss. nach dem finnischen Zikadologen Paavo Kontkanen (1905–1976) benannt kozhevnikovi Zachv. nach dem russischen Entomologen und ehemaligen Direktor des Moskauer Zoologischen Museums Gregorij Alexandrovich Kozhevnikov (1866–1933) kratochvili Lang nach dem tschechischen Entomologen Josef Kratochvil (1909–1992) benannt krueperi Fieb. nach dem deutschstämmigen, aber vorwiegend in Athen lebenden Ornithologen und Entomologen Theobald Johannes Krüper (1829–1917) benannt lacteinervis Kbm. der milchadrige (lat.) laeta H.-S. die fröhliche, heitere (lat.) laetifica Metc. die erheiternde (lat.) laevis Rib. der zarte (lat.) lamellaris Rib. mit einer Lamelle versehen (lat.) laminatus Fl. mit einem Blatt versehen (lat.), bezogen auf die Fühlerpalette der Männchen languidus Fl. der schwache (lat.) lanio L. der Fleischer (lat.), bezogen auf die manchmal rötliche Färbung des Vorderkörpers lanternae W. Wg. aus der Laterne (lat.), bezogen auf den Fundort des Typusmaterials

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 299 H. NICKEL lanzkei W. Wg. nach dem Sammler A. Lanzke aus Engern an der Weser lapidicola Vid. der Steinbewohner (lat.) lapponica Zett. die lappländische (nlat.) larvatus H.-S. der besessene (lat.) lasiocarpae Oss. der Faden-Segge (Carex lasiocarpa) zugehörig latifrons Kbm. die breite Stirn (lat.) latinus Lnv. der aus Latium stammende (lat.) lauri Horv. dem Lorbeer zugehörig (lat.) lelievrei Leth. nach dem französischen Lepidopterologen Ernest Lelièvre (1826–1889?) benannt lepidus Boh. der niedliche, anmutige (lat.) leporinus L. von lepus (lat.: Hase), bezogen auf die haarbüscheligen Wachsausscheidungen leptosoma Fl. der kleine Körper (griech.) leptus Fieb. der kleine, schmale (lat.) lequesnei W. Wg. nach dem Jerseyer Entomologen und Chemiker Walter John Le Quesne (1922–2006) benannt lethierryi Edw. nach dem französischen Hemipterologen Lucien François Lethierry (1830–1894) benannt leucophaea Preyssl. die weißgraue (griech.) ligustinus W. Wg. der ligurische (lat.) limbatellus Zett. der etwas gesäumte (lat.) limbatus Sign. der gesäumte (lat.) limicola Edw. der Schlammbewohner (lat.) limpidus W. Wg. der helle (lat.) lindbergi Lnv. nach dem finnischen Hemipterologen Håkan Lindberg (1898–1966) benannt lineatopunctata Mats. die mit einer Punktreihe versehene (lat.) lineatus L. der mit einer Linie versehene, auch: der in Leinwand gekleidete (lat.) lineolatus Br. der schwach linierte (lat.) lithuanica Vilb. die litauische (nlat.) litoralis Reuter dem Strand zugehörig (lat.) littoralis Kuntze wie oben lituratus Fall. der beschmierte (lat.) livens Zett. der bleifarbene (lat.) lividus Edw. der bleifarbige (lat.) loewi Leth. nach dem Wiener Hemipterologen Paul Löw benannt longicauda Rem. der lange Schwanz (lat.) longiceps Kbm. der lange Kopf (lat.) longipennis Curt. mit langen Flügeln (lat.) longispinus W. Wg. der lange Stachel (lat.) longiventris J. Shlb. mit langem Bauch versehen (lat.)

300 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen lorifer Rib. der eine Peitsche tragende (lat.) ludus Dav. et De L. unbekannt, eventuell der Spieler (lat.) lugens Horv. der trauernde (lat.) lugubrina Boh. die trauernde (lat.) lugubris Sign. der trauernde (lat.) lunaris M. et R. die halbmondförmige (lat.) lunulatus Zett. der mit kleinen Mondflecken versehene (lat.) lurida Fieb. die blassgelbe (lat.) luteipes Fieb. der (orange-)gelbfüßige (lat.) macchiae Lnv. der Macchia zugehörig (nlat.) macrocephala Fieb. die großköpfige (griech.) mactata Germ. von mactatio (lat.: Gemetzel), nach der blutroten Färbung maculatus Rib. der gefleckte (lat.) maculiceps Boh. der gefleckte Kopf (lat.) maculosus Then der gefleckte (lat.) magnus Mity. der große (lat.) major Uhl. die größere (lat.) makarovi Zachv. nach dem russischen Naturschützer Vasilij Nikitich Makarov (1887–1953) benannt maligna Walsh die schädliche (lat.) manderstjernii Kbm. nach dem russischen Koleopterologen und General Alexander Graf von Manderstjerna (1817–1888) benannt marginata H.-S. die berandete (lat.) margineguttata Leth. die am Rand betropfte (lat.) marginicollis Spin. mit berandetem Hals (lat.) maritimus Perr. der zum Meer gehörige (lat.) marmoratus Fl. der marmorierte (lat.) maroccana Mel. die marokkanische (lat.) martigniaca Cer. die aus Martigny (Wallis) stammende (nlat.) mayri Fieb. nach dem österreichischen Hymenopterologen und Hemi- pterologen Gustav L. Mayr (1830–1908) benannt mediterranea Fieb. die am Mittelmeer lebende (lat.) medius M. et R. der mittlere (lat.) megerlei Fieb. nach Johann Carl Megerle von Mühlfeld (1765–1840), dem früheren Kustos des Naturhistorischen Museums Wien benannt melanocephala Fieb. die schwarzköpfige (griech.) melanochaetus Fieb. der mit schwarzer Mähne versehene (griech.) melanops Fieb. das schwarze Gesicht (griech.) melanopsis Hd. wie oben melichari Horv. nach dem tschechischen Arzt und Zikadologen Leopold Melichar (1856–1924) benannt melissae Curt. der Melisse zugehörig (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 301 H. NICKEL mendax Rib. der Lügner (lat.) mesomelas Boh. der in der Mitte schwarze (griech.) mesopyrrhus Kbm. der in der Mitte feuerfarbene (griech.) metrius Fl. der mäßige, geringe (griech.) micantula Zett. die kleine schimmernde (lat.) michalki W. Wg. nach dem Leipziger Heteropterologen Otto Michalk (1886–1966) benannt microcera Vilb. die kleinhörnige (griech.) minima J. Shlb. die kleine (lat.) minkii Kbm. nach dem Krefelder Koleopterologen und Hymenoptero- logen Wilhelm Mink (1807–1883) benannt minor Kbm. der kleinere (lat.) minuscula Horv. die ein bisschen kleine (lat.) minutus F. der winzige (lat.) mirabilis Orosz die wundersame (lat.) mixtus F. der gemischte (lat.), Bezug unklar mocsaryi Horv. nach dem ungarischen Entomologen (v.a. Hymenoptero- logen) Sandor Mocsáry (1841–1915) benannt modestus Scott der bescheidene (lat.), Bezug unklar moestus Boh. der traurige (lat.) mollicula Boh. die recht zarte, zärtliche (lat.) monoceros Rib. einhörnig (griech.) montana Scop. die den Bergen zugehörige (lat.) montandoni Put. nach dem französisch-rumänischen Hemipterologen Arnold Lucien Montandon (1852–1922) benannt monticola Rem. der Bergbewohner (lat.) morbillosus Mel. der kränkelnde (lat.) morio Fieb. der Hofnarr (lat.) mucronatus Rib. der stachelspitzige (lat.), bezogen auf die Form der Analrohranhänge mulsanti Fieb. nach dem französischen Entomologen und Ornithologen Martial Étienne Mulsant (1797–1880) benannt multinotatus Boh. der vielfach gekennzeichnete (lat.) multireticulata M. et R. die vielfach genetzte (lat.) muscaeformis Schrk. der mückenförmige (lat.) musivus Germ. der mosaikartige (lat.) mutilla Rib. die verstümmelte (lat.) najas Nast griechische Wassernymphe, bezogen auf den Lebensraum am Wasser nanus H.-S. der Zwerg (lat.) nardeti Rem. dem Borstgrasrasen zugehörig (nlat.) nassatus Germ. der mit einer Schlinge versehene (lat.), Bezug unklar nasutus H.-S. der großnasige (lat.)

302 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen nebulosa Ball die trübe, dunkle (lat.) neglecta W. Wg. die übersehene (lat.) nervosus L. der geaderte (lat.) nicolasi Leth. nach dem französischen Entomologen Hector-Ulysse Nicolas (1834–1899) nicolovae Dlab. nach der Sammlerin des Typusmaterials, Veselina Nikolova, benannt nigra Goeze die schwarze (lat.) nigricans Kbm. der schwärzliche (lat.) nigricornis J. Shlb. mit schwarzem Horn (lat.), bezogen auf die Antennenbasen nigrifrons Kbm. der schwarzstirnige (lat.) nigriloba Edw. die schwarzlappige (lat./griech.) nigrita Kbm. die geschwärzte (lat.) nigritarsis Rem. mit schwarzem Tarsus (lat./griech.) nigrolineata Scott die schwarz gestreifte (lat.) nigropicta W. Wg. die schwarz bemalte (lat.) nisamiana Logv. nach der Region bzw. dem Ort Nizami in Aserbaidschan nitidissimus H.-S. der am meisten glänzende (lat.) nitidulus F. der etwas glänzende (lat.) nivea M. et R. die schneeweiße (lat.) nobilis Fieb. der vornehme (lat.) nodosus Rib. der knotige (lat.), bezogen auf die knotige Ausbuchtung des Aedeagus notata Curt. die gekennzeichnete (lat.) notula Germ. das Pünktchen (lat.) novaki W. Wg. nach dem dalmatinischen Entomologen Petar Novak (1879–1968) benannt nubilus Oss. der wolkige (lat.) obliqua W. Wg. die schräge (lat.) obscurella Boh. die etwas finstere (lat.) obsoletus Germ. der abgenutzte, schäbige (lat.) obtusa Vid. die stumpfe (lat.) obtusiceps Kbm. der stumpfköpfige (lat.) obtusifrons Stål die stumpfe Stirn (lat.) obtusivalvis Kbm. mit versteckten Klappen (lat.) occidentalis die westliche (lat.) Huang et Bourg. occirrega Rem. et Gugl. kombiniert aus occidentalis (lat.: die westliche) und irregulata (siehe dort) ocellaris Fall. mit Äuglein (lat.), bezogen auf die Ozellen oder ozellenartige Flecken ocellata Prov. die mit kleinen Augen versehene (lat.) ochreata Vilb. die mit einem Schienbeinschützer versehene (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 303 H. NICKEL ochrifrons Vilb. die ockerfarbene Stirn (lat.) ocularis M. et R. frei: mit Augenflecken (lat.) oculatus Ldb. wie oben odryssius Dlab. der Thraker (lat.) ognevi Zachv. nach dem russischen Theriologen Sergeij Ivanovich Ognev (1886–1951) benannt ogumae Mats. nach dem japanischen Sammler M. Oguma benannt ohausi W. Wg. nach dem Mainzer Koleopterologen Friedrich Ohaus (1864–1946) benannt okaensis Zachv. zum russischen Fluss Oka gehörig olitoria Say die dem Gemüse zugehörige (lat.) onustus Fieb. der beladene (lat.) ordinaria Rib. die ordentliche, vorzügliche (lat.) orichalceus Thoms. der messingfarbene (lat.) orientalis An. der östliche, auch: der orientalische (lat.) origani Zachv. dem Oregano zugehörig (lat.) ornata Mel. die geschmückte (lat.) ornaticeps Horv. mit geschmücktem Kopf (lat.) ornatus Perr. der geschmückte (lat.) orni L. der Manna-Esche (Fraxinus ornus, lat.: ornus) zugehörig orthocephalus Ferr. der mit aufrechtem Kopf (griech.) oshanini Razv. nach dem russischen Hemipterologen Vasilij Fedorovich Oshanin (1844–1917) benannt ossiannilssoni Ldb. nach dem schwedischen Hemipterologen Frej Ossiannilsson (1908–1995) benannt oxyura Hpt. der Spitzschwanz (griech.) pallens Stål die blasse (lat.) palliceps Horv. die blassköpfige (lat.) pallidinervis Dhlb. der bleichnervige (lat.) pallidula Boh. die ziemlich blasse (lat.) pallidus Dlab. der blasse (lat.) pallifrons Horv. die blassstirnige (lat.) pallipes Fieb. der blassfüßige (lat.) paludosa Boh. die sumpfige (lat.) palustris Rib. dem Sumpf zugehörig (lat.) pannonica Mats. die pannonische (lat.) panzeri Fl. nach dem bayerischen Entomologen und Botaniker Georg Wolfgang Franz Panzer (1755–1829) benannt paolii Oss. nach dem italienischen Akarologen und Entomologen Guido Paoli (1881–1947) benannt paradoxum H.-S. das seltsame (griech.) parcanicus Dlab. nach dem südslowakischen locus typicus Parkan (heute Štúrovo) benannt

304 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen parnassia Stål vom Parnass-Gebirge (Griechenland) stammend parvicauda Rib. der kurze Schwanz (lat.) parvispinus W. Wg. der kleine Dorn (lat.) parvula Boh. die sehr kleine (lat.) paryphasma Fl. die verwandte Gestalt (griech.) pascuellus Fall. auf der Weide (lat.) pascuorum Rib. den Weiden zugehörig (lat.) pauxillus Fieb. der ganz kleine (lat.) pectoralis Mats. auf die Brust bezogen (lat.) pedemontana Vid. die am Fuß der Berge lebende, hier auch: die piemontesische (lat.) pellax Horv. der listenreiche, verführerische (lat.) pellucida F. die durchscheinende (lat.) penthopitta Walk. frei: der pechschwarze; von penthos (griech.: Trauer) und pitta (griech.: Pech) perplexus Rib. der verworrene (lat.) perrieri Rib. nach dem französischen Zoologen Rémy Perrier (1861– 1936) benannt persimilis Edw. die sehr ähnliche (lat.) perspicillata Boh. die durchsichtige (lat.) petryi Schuhm. nach dem Thüringer Entomologen und Botaniker Arthur Petry (1858–1932) benannt philesakis Hoch die Freundin von Sakis (griech.), bezogen auf den griechischen Zikadologen Sakis Drosopoulos (* 1944) phragmitis Boh. dem Schilfrohr zugehörig (lat.) pictilis Stål die bemalte (lat.) pictipennis Kbm. der buntflügelige (lat.) picturatus C. Shlb. der gestickte, bunt bemalte (lat.) pictus Leth. der (bunt) bemalte (lat.) pilatoi D’Urso et Gugl. nach dem sizilianischen Zoologen Giovanni Pilato benannt pilosus Ol. der behaarte (lat.) placida Horv. die sanfte, friedliche, ruhige (lat.) planicollis Thoms. der breithalsige (lat.) planiscuta Thoms die mit breitem Schild versehene (lat.) platanicola Vid. der Platanenbewohner (lat.) plebejus Scop. der gemeine (lat.) plurispinosa W. Wg. die vieldornige (lat.) plutonius Uhl. dem Pluto (römischer Gott der Unterwelt) zugehörig, bezogen auf die schwärzliche Färbung podolica Eichw. die podolische (lat.) poecilus H.-S. der buntgefleckte (griech.) populi L. der Pappel zugehörig (lat.) porcus Em. das Schwein (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 305 H. NICKEL potentillae Mor. dem Fingerkraut zugehörig (lat.) praecox Hpt. die frühzeitige (lat.) prasina Boh. die lauchgrüne (lat.) prasinula Fieb. die etwas lauchgrüne (lat.) preyssleri H.-S. nach dem böhmischen Entomologen Jan Daniel Preyssler (1768–1839) benannt proceps Kbm. der vorstehende Kopf (lat.) procerus H.-S. der schlank gewachsene (lat.) prolixa Leth. die verlängerte (lat.) propinqua Fieb. die Verwandte (lat.) provincialis Rib. der provenzalische (lat.) pruinosa Say die bereifte (lat.) pruni Rib. der Schlehe (wörtlich: der Pflaume) zugehörig (lat.) prunicola Edw. der Pflaumenbewohner (lat.) pseudocellaris Fl. mit falschen Äuglein (griech./lat.), bezogen auf ozellen- artige Flecken pseudomicantula Knight die falsche Micantula (siehe dort; nlat.) pteridis Spin. dem Farn zugehörig (griech.) pulchellus Curt. der kleine hübsche (lat.) pulchra P. Löw die herrliche (lat.) pulchripennis Asche mit schönen Flügeln (lat.) pulicaris Fall. der flohartige (lat.) pullula Boh. frei: die sehr kleine niedliche (lat.) punctata F. die punktierte (lat.) puncticollis H.-S. mit gepunktetem Hals punctifrons Fall. die punktierte Stirn (lat.) punctulum Kbm. das Pünktchen (lat.) punctum Fl. der Punkt (lat.) pungens Rib. die stechende (lat.), nach den zugespitzten Anal- rohranhängen pusilla Mats. die winzige (lat.) putoni Then nach dem Lothringer Hemipterologen Jean Baptiste Auguste Puton (1834–1913) benannt pygmaeus Vilb. der zwergenhafte (lat.) pyrenaea Fieb. die pyrenäische (griech./lat.) quadratus Forel der viereckige (lat.) quadricornis Dlab. der mit vier Hörnern versehene (lat.) quadrimaculatus Sign. der vierfleckige (lat.) quadrinotata F. die vierfach gekennzeichnete (lat.) quadripunctulatus Kbm. der mit vier Pünktchen versehene (lat.) quadrisignata Hag. die vierfach gezeichnete (lat.) quadrivittatus Fieb. der mit vier Binden geschmückte (lat.) quadrum Boh. das Viereck, der Quader (lat.)

306 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen quercus F. die Eiche (lat.) quinquecostatus Duf. der mit fünf Rippen versehene (lat.) quinquenotata Boh. die fünffach gekennzeichnete (lat.) radiatus O. Costa der mit Strahlen umgebene (lat.) raniformis M. et R. der froschförmige (lat.) remanei Nick. nach dem deutschen Hemipterologen Reinhard Remane (1929–2009) benannt remotus Edw. der entfernte (lat.) renonensis W. Wg. nach der Südtiroler Gebirgsregion Ritten (ital.: Renon) benannt repletus Fieb. der ergänzte (lat.) reticulata F. frei: die netzartige (lat.) rex Gugl. et Bück. von Tyrannosaurus rex übernommen reyi Fieb. nach dem französischen Entomologen Claudius Rey (1817–1895) benannt rhamni Ferr. zum Kreuzdorn (Rhamnus frangula) gehörend (lat.) rhamnicola Horv. der Bewohner von Kreuzdorn (Rhamnus frangula) (nlat.) rhenana W. Wg. die Rheinische (lat.) rhodophila Cer. die rosenliebende (griech.) rhombifer Fieb. der Rautenträger (griech./lat.), bezogen auf die Scheitel- zeichnung ribauti Oss. nach dem französischen Hemipterologen und Myriapodo- logen Henri Ribaut (1872–1967) benannt rorida M. et R. die mit Tau benetzte (lat.) rosae L. der Rose zugehörig (lat.) rosaesugans Cer. die an Rosen saugende (lat.) rosaurus Gugl. et Bück. von Rosetta (italienischer Vorname) und sauros (griech.: Echse, Drache) rosea Fl. die rosenfarbige (lat.) roseipennis Tollin mit rosenfarbenen Flügeln (lat.) rosincola Cer. der Rosenbewohner (lat.) rostrata Rib. die geschnäbelte (lat.) rotundiceps Leth. der runde Kopf (lat.) rotundifrons Kbm. der rundstirnige (lat.) rubi Boh. zur Himbeere (oder Brombeere) gehörig (lat.) rubroflava Lnv. die rotgelbe (lat.) rubrovittata Leth. die rotstreifige (lat.) rufescens Mel. der rötlich werdende (lat.) ruficeps Mel. die rotköpfige (lat.) russeola Fall. die rötliche (lat.) ruthenica Zachv. die ruthenische (lat.) rutilans Kbm. der rötlich schimmernde (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 307 H. NICKEL saageri W. Wg. nach Hans Saager (1887–1980) benannt, einem ehrenamtlichen Mitarbeiter des Lübecker Naturkunde- museums (Heteroptera, Hymenoptera, Diptera) sabaudiana Lall. die aus Savoyen stammende (lat.) sabulicola Curt. der Sandbewohner (lat.) sachalinensis Mats. der aus Sachalin stammende (nlat.) sagittarius Rib. der Bogenschütze (lat.), bezogen auf die Form der Penis- spitze sahlbergi Fl. nach dem finnischen Zikadologen und Koleopterologen Johan Reinhold Sahlberg (1845–1920) benannt salicicola Edw. der Weidenbewohner (lat.) salicina Goeze den Weiden zugehörig (lat.) salina Hpt. die salzige (lat.) salsolae Put. dem Salzkraut zugehörig (nlat.) saltuella Kbm. auf der Waldwiese (lat.) salviae Arz. et Vid. dem Salbei zugehörig (lat.) sanguinolenta Scop. die blutbefleckte (lat.) sanguinosa R. die blutrote (lat.) sardus Rib. der Sarde (lat.) saturata Edw. die reichlich gefärbte (lat.) saurosus Gugl. et Bück. Anagramm von rosaurus (siehe dort) scabripennis Edw. der rauhflügelige (lat.) scalaris Rib. die mit Leitersprossen versehene (lat.) scanicus Fall. die skanische (lat.), Bezug zur südschwedischen Region Skane schaefferi Metc. nach dem Regensburger Entomologen Gottlieb August Wilhelm Herrich-Schäffer (1799–1874) benannt schenckii Kbm. nach dem hessischen Hymenopterologen Philipp Adolph Schenck (1803–1878) benannt schmidtgeni W. Wg. nach dem ehemaligen Direktor des Naturhistorischen Museums Mainz, Otto Schmidtgen (1879–1938), benannt schmidti W. Wg. nach dem Fürther Heteropterologen und Medizinalrat Karl Schmidt (1892–1940) benannt schneideri Günth. nach dem Schweizer Entomologen Otto Schneider-Orelli (1880–1965) benannt scotti Edw. nach dem englischen Entomologen John Scott (1823–1888) benannt scurra Germ. der Stutzer; aber auch: Possenreißer (lat.) scutellaris H.-S. mit auffälligem Schildchen versehen (lat.) scutellata Boh. wie oben scutuliferus W. Wg. der ein kleines Schild trägt (lat.) sellatus Uhl. der mit einem Sattel versehene (lat.), bezogen auf die Flügelzeichnung

308 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen semipunctata Fieb. die halb gepunktete (lat.) septemnotatus Fall. der siebenfach gekennzeichnete (lat.) septentrionalis W. Wg. die nördliche (lat.), wörtlich: zum Sternbild Septentriones (= Großer Wagen) gehörend serrata Vilb. die gesägte (lat.) serratulae F. im Sinne von Fabricius: zur Acker-Kratzdistel (Cirsium arvense, früherer Name: Serratula arvensis) gehörig (lat.) servadeii Vid. nach dem italienischen Entomologen Antonio Servadei (1908–1979) benannt severtsovi Zachv. nach dem russischen Zoologen Nikolai Alexejewitsch Severtsov (1827–1885) benannt sexmaculata Hd. die sechsfach gefleckte (lat.) sexnotatus Fall. der sechsfach gekennzeichnete sibiricus Horv. der sibirische (lat.) sicula Mats. die sizilianische (lat.) signatipennis Boh. mit gezeichneten Flügeln (lat.) signifer Then der Zeichenträger (lat.) silvicola Oss. der Waldbewohner (lat.) sima Rib. die stumpfnasige (lat.) similis Kbm. die ähnliche (lat.) simillima W. Wg. die sehr ähnliche (lat.) simplex H.-S. der einfache (lat.) singeri W. Wg. nach dem Aschaffenburger Arzt und Entomologen Karl Singer (1865–1959) benannt sinuata Then die gebogene (lat.), bezogen auf den Hinterrand des 7. Abdominalsternites der ♀ slovacus Dlab. der slowakische (lat.) smaragdinus Em. der smaragdgrüne (lat.) smaragdula Fall. kleiner Smaragd (lat.), nach der Färbung smreczynskii Dwor. nach dem Krakauer Entwicklungsbiologen und Koleoptero- logen Stanislaw Smreczynski benannt (1899–1975) benannt sociabilis Oss. die gesellige (lat.) socialis Fl. der kameradschaftliche (lat.) solani Curt. zum Nachtschatten gehörig (lat.), gemeint ist hier die Kartoffel soosi Orosz nach dem ungarischen Entomologen und Zoologen Árpád Soós (1912–1991) benannt sorbi W. Wg. der Mehlbeere zugehörig (lat.) sordidipennis Stål mit schmutzigen Flügeln (lat.) sordidula Stål die etwas schmutzige, armselige (lat.) sordidus Zett. der schmutzige (lat.) soror Lnv. die Schwester (lat.) sororcula Horv. die kleine Schwester (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 309 H. NICKEL spathulata Rib. die mit einem Spatel versehene (lat.) speciosa Boh. die ansehnliche (lat.) sphagneticus Em. der dem Torfmoos zugehörige (lat.) spinigera Edw. die dornentragende (lat.) spinosa Fieb. die dornige (lat.) splendidulus Fieb. der etwas glänzende (lat.) spoliata Horv. die geplünderte (lat.) spumarius L. der schaumige (lat.) stachydearum Hd. den Ziesten zugehörig (lat.) stactogalus Fieb. frei: der mit Milch betröpfelte (griech.) stali Metc. nach dem schwedischen Hemipterologen und Koleoptero- logen Carl Stål (1833–1878) benannt staminata Rib. die mit Faden versehene (lat.), bezogen auf die sehr dünnen Aedeagusanhänge staurus Iv. von stauros (griech.: Pfahl) stehliki Laut. nach dem tschechischen Heteropterologen Jaroslav L. Stehlík (*1923) benannt steini Fieb. vermutlich nach dem Berliner Sammler Friedrich Stein benannt stellulata Burm. mit Sternchen übersät (lat.) stenoptera Fl. die schmalflügelige (griech.) steveni Krynicki nach dem schwedisch-russischen Botaniker und Entomologen Christian Christianowitsch Steven (1781– 1863) benannt sticticus R. der punktierte (griech.) stigmaticalis Lew. der gebrandmarkte (lat.) stigmaticus Germ. wie oben stigmatipennis M. et R. mit gebrandmarkten Flügeln (lat.) straminea Stål die strohfarbene (lat.) striatus L. der gestreifte (lat.) striatella Fall. die fein gestreifte (lat.) strigilifer Oss. Striegel tragend (lat.), nach der Form des Penis striifrons Kbm. gestreifte Stirn (lat.) striola Fall. der Streif (lat.) strobli W. Wg. nach dem steirischen Naturforscher (v.a. Dipterologen) Gabriel Strobl (1846–1925) benannt stylata Boh. die mit einer Stange versehene (griech.), bezogen auf den langen Ovipositor suavis R. die liebliche (lat.) subangulata J. Shlb. die schwach gewinkelte (lat.) subfusculus Fall. die etwas bräunliche (lat.) subquadrata H.-S. die fast viereckige (lat.) substriatus Then kleiner als striatus (siehe dort; lat.)

310 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen subulata Rib. die mit einer (pfriemenartigen) Spitze versehene (lat.) sudeticus Kol. der sudetische (nlat.) sulcata Rib. die gefurchte (lat.) sulphurea M. et R. die schwefelgelbe (lat.) sulphurella Zett. die etwas schwefelfarbene (lat.) sursumflexus Then der aufwärts gebogene (lat.) taeniatifrons Kbm. die mit Band versehene Stirn (lat.) taeniatus Horv. der gebänderte (lat.) tamaninii Lnv. nach dem italienischen Hemipterologen Livio Tamanini (1907–1997) benannt tamaricis Put. der Tamariske zugehörig (lat.) tapinus Fieb. der niedere (griech.) tarda Hpt. die langsame (lat.) taunica W. Wg. die aus dem Taunus stammende (lat.) taurus Rib. der Stier (lat.), nach der Form des Aedeagus temperei Rib. nach dem französischen Koleopterologen Gaston Tempère (1900–1985) benannt tenella Fall. die sehr zarte (lat.) tenerrima H.-S. wie oben: die sehr zarte (lat.) tenuis Germ. die schmale (lat.) tersa Edw. die reine (lat.) tessellata Leth. die gewürfelte, schachbrettartige (lat.) teucrii Cer. dem Gamander zugehörig (lat.) thenii P. Löw nach dem österreichischen Zikadologen Franz Then (1841–1919) benannt thoulessi Edw. nach dem englischen Entomologen Henry James Thouless (1864–1933) benannt tiaratus Fieb. der mit einer Tiara versehene (pers./griech./lat.) tibialis Panz. die mit besonderer Tibia versehene (lat.) tiliae Fall. der Linde zugehörig (lat.) titanus Ball der riesige (lat.) tithide Ferr. unklar, eventuell von tithi (sanskr.: lunarer Tag; oder auch 15) topoli Zachv. der Pappel (russ.: topol) zugehörig torneellus Zett. nach der Herkunft: Lapponia tornensis (nördlicher Teil Lapplands) translucidus M. et R. der durchscheinende (lat.) transsylvanica Fieb. die transsylvanische (lat.) transversa Fall. die quer liegende (lat.), bezogen auf das Scheitelband tremulae Estl. der Zitter-Pappel (Populus tremula) zugehörig tricinctus Curt. der drei Gürtel tragende (lat.) tricolor O. Costa der dreifarbige (lat.) trifasciata Fourcr. die dreifach gebänderte (lat.)

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 311 H. NICKEL tristis Zett. die traurige (lat.) trivia Germ. vermutlich: die abgeriebene (lat.) truncatus Fieb. der gestutzte (lat.) tshinari Zachv. von tshinar (russ.: Platane) tyrannus Gugl. et Bück. von Tyrannosaurus rex übernommen ukrainicus Logv. der ukrainische (nlat.) ulicis Rib. dem Stechginster (Ulex) zugehörig (lat.) ulmi L. der Ulme zugehörig (lat.) ulmiphagus Wls. et Clr. der Ulmenfresser (lat./griech.) unciger Rib. der Hakenträger (lat.) undatus De G. der gewellte (lat.) unicolor H.-S. die einfarbige (lat.) untica Dlab. nach dem nord-mongolischen locus typicus Unt benannt urticae F. der Brennnessel zugehörig (lat.) usnus Beirne unklar, keine derivatio nominis ustulatus M. et R. der etwas gebräunte (lat.) variatus Fall. der gesprenkelte (lat.) variegatus Kbm. der bunt gemachte (lat.) vasconica Rib. die baskische (lat.) velata Rib. die mit Segel versehene (lat.) veneta Dlab. die venetische (lat.) venosus Germ. der geaderte (lat.) ventouxiana Biem. die vom Mont Ventoux stammende (nlat.) ventralis Fall. mit auffälligem Bauch versehen (lat.) venustulus Germ. der etwas anmutige (lat.) verbae Zachv. zur Weide (russ.: verba) gehörig versuta Mel. die täuschende (lat.) vestita Rib. die bekleidete (lat.) viaduensis W. Wg. zum Fluss Oder (lat.: Viadua) gehörig vicinus Mel. der benachbarte (lat.) vinealis Biem. wörtlich: zum Weinberg gehörig (lat.), hier jedoch bezogen auf die Wirtspflanze Agrostis vinealis virescens Panz. grün werdend (lat.) virgator Rib. der mit Ruten schlagende (lat.), bezogen auf die langen, verschmälerten Analrohranhänge viridigriseus Edw. der grüngraue (lat.) viridinervis W. Wg. die grünadrige (lat.) viridis L. die grüne (lat.) vitis Göthe der Weinrebe zugehörig (lat.) vitreus F. der gläserne (lat.) vitripennis Fl. mit gläsernen Flügeln (lat.) vittata L. die mit einer Binde geschmückte (lat.) vittifrons Kbm. gestreifte Stirn (lat.) vittipennis J. Shlb. mit gestreiften Flügeln (lat.)

312 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Etymologie der Zikadennamen volgensis Prid. von der Wolga stammend (lat.) vulnerata Rossi die verwundete (lat.) wagneri China nach dem Hamburger Lehrer und Hemipterologen Wilhelm Wagner (1895–1977) benannt wagnerianus Nast siehe oben wahlbergi Boh. nach dem schwedischen Naturforscher Johan August Wahlberg (1810–1859) benannt wallengreni Stål nach dem schwedischen Lepidopterologen Hans Daniel Johan Wallengren (1823–1894) benannt wilhelmi An. et Av. nach dem Hamburger Lehrer und Hemipterologen Wilhelm Wagner (1895–1977) benannt williamsi China nach dem britischen Sammler B.S. Williams (1891–1941?) benannt xantha Vilb. die gelbe, blonde (griech.) xanthoneurus Fieb. der gelbnervige (griech.) yarkonensis Lnv. vom Fluss Yarkon in Israel stammend zachvatkini Em. nach dem russischen Hemipterologen und Akarologen Alexej Alexejewitsch Zachvatkin (1906–1950) benannt zaisanica Mity. nach der im östlichen Kasachstan gelegenen Region Zaisan (nlat.) zelleri Kbm. nach dem deutschen Mikrolepidopterologen Philipp Christoph Zeller (1808–1883) benannt zonata Em. die mit einem Gürtel versehene (lat.)

Danksagung Diese Arbeit ist meinem lieben Kollegen und lange Zeit väterlichen Freund Pavel Lauterer in dankbarer Verbundenheit gewidmet. Für Hinweise und Diskussionen danke ich Georgii Anufriev (Nizhny Novgorod), Michel Boulard (Paris), Christoph Bückle (Tübingen), André Burgers (Minas de Riotinto), Henry Callot (Strasbourg), Dmitri Dmitriev (Champaign), Irena Dworakowska (Vancouver), Sasha Emeljanov, Vladimir Gnezdilov (beide St. Petersburg), Ernst Heiss (Innsbruck), Günter Hofmann (Stockstadt), Yoshizawa Kazunori (Sapporo), Wolf-Rüdiger Klingler (Rheinbach), Igor Malenovský (Brno), Merv Nielson (Provo), Francesco Poggi (Missaglia), Matthias Schaefer, Michael Schmidt (beide Göttingen), Adéline Soulier-Perkins (Paris) und Chandra Viraktamath (Bangalore).

Zusammenfassung Diese Arbeit bietet Übersetzungen der wissenschaftlichen Namen der Gattungen, Untergattungen, Arten und Unterarten der in Mittel- und Nordeuropa vorkommenden Zikaden (Hemiptera: Auchenorrhyncha), einschließlich einiger Synonyme. Insgesamt werden 363 supraspezifische und 946 spezifische und subspezifische Namen behandelt. In einigen weniger ersichtlichen Fällen werden auch Anmerkungen zur Interpretation geliefert.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 313 H. NICKEL

Literatur

ANONYMUS 2013a: Pollux: Archimedes Project Dictionary Access. http://archimedes.fas.harvard.edu/pollux (Zugriff Juni bis August 2013). ANONYMUS 2013b: Wikipedia, die freie Enzyklopädie. http://de.wikipedia.org (Zugriff Juni bis August 2013). ANONYMUS 2013c: Wikipedia, the free encyclopedia. http://en.wikipedia.org (Zugriff Juni bis August 2013). ANONYMUS 2013d: ZoBoDat: Zoologisch-Botanische Datenbank des Biologiezentrums Linz der Oberösterreichischen Landesmuseen. Biografien von Biologen und Geowissenschaftern. http://www.zobodat.at (Zugriff Juni bis August 2013). ASPÖCK H. & ASPÖCK U. 2013: Woher kommen die Namen? Die validen rezenten Taxa der Kamelhalsfliegen der Erde: Systematisches Verzeichnis und Etymologie (Insecta: Endopterygota: Neuropterida: Raphidioptera). Entomologica Austriaca 20: 9–155. BIEMAN K. DEN, BIEDERMANN R., NICKEL H. & NIEDRINGHAUS R. 2011: The Planthoppers and Leafhoppers of Benelux–Identification keys to all families and genera and all Benelux species not recorded from Germany. Cicadina, Suppl. 1: 1–120. BORODIN O. 2004: A checklist of the Auchenorrhyncha of Belarus (Hemiptera, Fulgoromorpha et Cicadomorpha). Beiträge zur Zikadenkunde 7: 29–47. BROWN R.W. 1954: Composition of scientific words. Published by the author, Baltimore. 885 pp. CARPENTER M. M. 1945: Bibliography of biographies of entomologists. The American Midland Naturalist 33(1):1–116. CARPENTER M. M. 1953: Bibliography of biographies of entomologists (Supplement). The American Midland Naturalist 50(2): 257–348 DURBEŠIÆ P. 2011: Croatian Entomofauna – Looking back from the present and future plans. Entomologia Croatica 15(1–4): 17–101. D’URSO V. 1995: Homoptera Auchenorrhyncha. In: MINELLI A., RUFFO S. & LA POSTA S. (eds.): Checklist delle specie della Fauna Italiana 42. Calderini, Bologna, 35 pp. EMMRICH R. 2013: Geschichte der Zikadenkunde in Mitteleuropa. Pp. 19–25. In: HOLZINGER W.E., KAMMERLANDER I. & NICKEL H.: The Auchenorrhyncha of Central Europe. Die Zikaden Mitteleuropas. Volume 1: Fulgoromorpha, Cicadomorpha excl. Cicadellidae. Brill, Leiden – Boston, 673 pp. FLIEDNER H. 1997: Die Bedeutung der wissenschaftlichen Namen europäischer Libellen. Libellula, Supplement 1: 1–111. GEMOLL W. 1997: Griechisch- Deutsches Schul- und Handwörterbuch. 9. Auf. (Nachdruck). Hölder-Pichler- Tempsky, Wien. 860 pp. GENAUST H. 1996: Etymologische Wörterbuch der botanischen Pflanzennamen. 3. Aufl. Springer, Basel. 701 pp. GÜNTHART H. & MÜHLETHALER R. 2002: Provisorische Checklist der Zikaden der Schweiz (Insecta: Hemiptera, Auchenorrhyncha). Denisia 4: 329–338. GYÖRFFY G., KISS B., KOCZOR S. & OROSZ A. 2009: Checklist of the fauna of Hungary. Volume 4. Hemiptera: Archaeorrhyncha, Clypeorrhyncha. Hungarian Natural History Museum, Budapest, 79 pp. HENTSCHEL E. & WAGNER G. W. 1996: Zoologisches Wörterbuch. 6. Aufl. Spektrum, Heidelberg. 680 pp. HORN W. & SCHENKLING S. 1928–1929: Index Litteraturae Entomologica. Serie I: Die Welt-Literatur über die gesamte Entomologie bis inklusive 1863. Berlin-Dahlem, Selbstverlag W. Horn, 1–4, 1426 pp. HOLZINGER W.E. 2009: Auchenorrhyncha (Insecta). Checklisten der Fauna Österreichs 4. Biosystematics and Ecology Series 26: 41-100. HOLZINGER W. E. & SELJAK G. 2001: New records of planthoppers and leafhoppers from Slovenia, with a checklist of hitherto recorded species (Hemiptera: Auchenorrhyncha). Acta Entomologica Slovenica 9: 39–66. KERZHNER I. M. 2007: The etymology of Fabrician generic names in Hemiptera. Zoosystematica Rossica 15 (2006): 304. MÜHLETHALER R., HOLLIER J., NICKEL H., GNEZDILOV V. M., WILSON M. R., KUNZ G., GÜNTHART H. 2009: Neue und bislang selten gesammelte Zikaden aus der Schweiz (Hemiptera, Auchenorrhyncha). Entomo Helvetica 2: 39–48. NICKEL H. 2010: First addendum to the Leafhoppers and Planthoppers of Germany (Hemiptera: Auchenorrhyncha). Cicadina 11: 107–122.

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NICKEL H. & REMANE R. 2002: Artenliste der Zikaden Deutschlands, mit Angaben zu Nährpflanzen, Nahrungsbreite, Lebenszyklen, Areal und Gefährdung (Hemiptera, Fulgoromorpha et Cicadomorpha). Beiträge zur Zikadenkunde 5: 27–64. ÖSTERREICHISCHE AKADEMIE DER WISSENSCHAFTEN 2013: Österreichisches Biographisches Lexikon 1815–1950, Online-Edition. http://www.biographien.ac.at (Zugriff Juli 2013). REDL M. & KALLENBORN H. 1995: Etymologie der Gattungsnamen mitteleuropäischer Wanzen (Insecta: Heteroptera). Mitteilungen des Internationalen Entomologischen Vereins, Supplement 3: 1–29. SCHENKLING S. 1917: Erklärung der wissenschaftlichen Käfernamen aus Reitter’s Fauna Germanica. Schriften des deutschen Lehrervereins für Naturkunde 34: 1–80. SÖDERMAN G., GILLERFORS G. & ENDRESTÖL A. 2009: An annotated catalogue of the Auchenorrhyncha of Northern Europe (Insecta, Hemiptera: Fulgoromorpha et Cicadomorpha). Cicadina 10: 33–69. STOWASSER J. M., PETSCHENIG M., SKUTSCH F. 1979: Lateinisch-Deutsches Schulwörterbuch. Hölder-Pichler- Tempsky, Wien. 574 pp. WERNER F. C. 1968: Wortelemente lateinisch-griechischer Fachausdrücke in den biologischen Wissenschaften. Suhrkamp Taschenbuch 64, Verlag Enzyklopädie, 3. Auflage, Leipzig, 475 pp. WISSENSCHAFTLICHER RAT DER DUDENREDAKTION 1997: Duden Fremdwörterbuch. Dudenverlag, Mannheim. 864 pp.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 317–325, 2013

Two new species of Oncylocotis (Hemiptera: Heteroptera: ) from Australia

PETR BAÒAØ1* & PAVEL ŠTYS2 1 Department of Entomology, Moravian Museum, Hviezdoslavova 29a, Brno, CZ-627 00, Czech Republic; e-mail: [email protected] 2 Department of Zoology, Faculty of Science, Charles University in Prague, Vinièná 7, CZ-128 44 Praha 2, Czech Republic; e-mail: [email protected] * Corresponding author

BAÒAØ P. & Š TYS P. 2013: Two new species of Oncylocotis (Hemiptera: Heteroptera: Enicocephalidae) from Australia. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 317–325. – Two new species of the genus Oncylocotis Stål, 1856: Oncylocotis stehliki sp.nov. and Oncylocotis lautereri sp.nov. from Northern Territory, Australia, are described and illustrated. Keywords. Heteroptera, Enicocephalomorpha, true bugs, Oncylocotis, taxonomy, morphology, Australasian Region, Northern Territory

Introduction Oncylocotis Stål, 1856 is the richest genus of the infraorder Enicocephalomorpha, with 100 described species and many hundreds undescribed, occurring in the Old World. However, the Australian fauna of the true bug infraorder Enicocephalomorpha is poorly known (see CASSIS & GROSS 1995). To date, three species of the genus Oncylocotis have been described from Australia. WESTWOOD (1837) described the first four species of enicocephalomorphans (in the genus Enicocephalus Westwood 1837, placed within family ; two of them actually belong to the genus Oncylocotis), one of them, O. tasmanicus (Westwood, 1837) from Tasmania. Since its description, this species has been collected several times in southern regions of Australia, and has recently become distributed in the south-east coastal area, New South Wales and Tasmania (CASSIS & GROSS, 1995). JEANNEL (1942) added two Australian species (both originally placed in the genus Didymocephalus Jeannel, 1942, junior synonym of Oncylocotis): Oncylocotis leoninus (Jeannel, 1942) from New South Wales and Oncylocotis saetosus (Jeannel, 1942) from Victoria. In this work we add two new species, the first members of this genus to be described from Northern Territory.

Material and methods By the term “dorsal ocular index” we indicate the ratio of the minimum width of the vertex to the maximum width of the eye; this is most easily measured as twice minimum interocular distance / (maximum width across eyes minus minimum interocular distance).

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Both the specimens studied are dry-mounted, with considerably deformed abdomens. Hence, abdomen and genitalia have not been studied in these unique examples. The label data are cited verbatim, including potential errors, using a slash (/) to separate rows on one label and double slash (//) for different labels. Colour photographs were taken with a Leica MSV266. Drawings and measurements were made using a SZP 11 ZOOM stereoscopic microscope with an eye-piece graticule. SEM photographs were provided by SEM JEOL 6380 LV. The type material is deposited in the collections of Museum and Art Gallery of the Northern Territory (MAGNT), Darwin, Northern Territory, Australia.

Taxonomy

Oncylocotis Stål, 1856 Type species: Oncylocotis nasutus Stål, 1856: 44; by monotypy.

Oncylocotis stehliki sp.nov. (Figs 1, 3, 5, 7–8, 11–12) Type material. Holotype: male, labelled: ‘AUSTRALIA, NT 37m alt / NITMILUK N.P. 3.12.2008 / 24°18′S, 131°34′E ; / EDIT FALLS, St. Jakl lgt.’ // ‘Holotype / Oncylocotis / stehliki sp. nov. / P. Baòaø & P. Štys det. 2013’ (MAGNT). Specimen dry-mounted on rectangular card, right fore leg mounted separately on the same card and gold-coated for SEM photography. Diagnosis. Taking into account the newly-described Oncylocotis stehliki sp.nov. and all the described and undescribed species of Oncylocotis of which we are aware, O. stehliki sp.nov. may be recognized by the following unique combination of characters: legs long and slender, hind lobe of pronotum very long and wide, conspicuous setigerous granules on head and pronotum, and short, conspicuously curved vestiture of dorsum of body. Description. Macropterous male (holotype). Measurements (in mm). Total body length – 5.33. Head (without neck). Total length – 0.91; posterior lobe, length – 0.38, posterior lobe, width – 0.42; distance of eye to apex of antennifer – 0.22; diatone (maximum width across eyes) – 0.49; dorsal synthlipsis (minimum interocular distance) – 0.24; eye, length – 0.21; gena, minimum width – 0.22; interocellar distance – 0.20; minimum distance ocellus–eye – 0.04. Labium. Total length – 0.92; segment I, length – 0.09; segment II, length – 0.11; segment III, length – 0.44; segment III, maximum height – 0.11; segment IV, length – 0.28. Antenna. Segment I, length – 0.18; segment II, length – 0.50; segment III, length – 0.42; segment IV, length – 0.51; segment II, basal width – 0.05, segment II, distal width – 0.06. Pronotum. Total length (maximum) – 0.87; collum, length (median) – 0.22, maximum width – 0.41; middle lobe, length (median) – 0.33, middle lobe, width (maximum) – 0.73; hind lobe, length (maximum) – 0.42, hind lobe, length (mediane) – 0.22, hind lobe, width (maximum) – 1.11. Fore leg. Femur, length – 0.89, femur, maximum width – 0.22; tibia, length – 0.80, tibia, maximum width – 0.20; tarsus, length – 0.18, tarsus, maximum width – 0.08; fore claws, length – 0.20. Fore wing. Maximum length – 3.20.

318 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Oncylocotis

Figs 1–6. 1, 3, 5 – Oncylocotis stehliki sp.nov., male holotype (5.33 mm); 2, 4, 6 – Oncylocotis lautereri sp.nov., female holotype (4.24 mm); 1–2 – dorsal habitus; 3–4 – head, lateral view; 5–6 – head, dorsal view.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 319 P. B AÒAØ & P. ŠTYS

Coloration. Body uniformly light brown (Fig. 1), anterior lobe of head conspicuously darker than posterior lobe. Legs, antennae and labium yellowish. Abdomen dark brown. Sculpture. All aspects of the head (except narrow median of anterior lobe, Fig. 7) and dorsal and dorsolateral faces of pronotum, except for pronotal constrictions and impressions on its middle lobe, with large setigerous tubercles. All body surfaces densely covered in very fine, minute, non-setigerous granules. Vestiture. Dorsal and dorsolateral surfaces of head with short, conspicuously bent, semi-erect to erect setae directed forwards; lateral surfaces with longer, semi-erect setae directed forwards; ventral face of preocular part (‘gena’) of anterior lobe with erect setae pointing in various directions; ventral part between eyes with long, erect setae directed backwards; venter of posterior lobe of head with long (longest on head) setae directed forwards. Labium densely covered in short, semi-erect setae, mixed with sparsely distributed, longer, more protruding ones. Antennae densely covered with short, semi- erect setae, on segments II–IV mixed with longer (longer as segment diameter), more protruding setae. Thorax and forewing veins with short, conspicuously curved, semi-erect to appressed setae, directed rearwards. Dorsal and anterodorsal faces of femora and tibiae of all pairs with short, strongly curved setae, ventral and posteroventral surfaces with long, dense, straight, erect setae, only apically curved. Structure. Head (Figs 3, 5, 7) slender, not quite as long as pronotum. Eyes large, prominent, ocular index 1.92. Posterior lobe with narrow median impression, slightly transverse, ratio of length to width 0.9. Ocelli very large, situated on small tubercles. Antennae long, only slightly shorter than head and pronotum together (ratio 0.9), antennal formula (longest segment first): IV=II-III-I. Labium (Fig. 3) long, reaching midway along posterior lobe, labial formula (longest segment first): III-IV-II-I. Thorax. Pronotum with very conspicuous constrictions. Collum long, ratio of length to width 0.54, with well-developed precollum. Middle lobe short and wide, more than twice as wide as long (ratio 2.21); with wide, inversely T-shaped impression in the middle, reaching from collum-middle lobe constriction almost to posterior margin of middle lobe. Sublateral parts of middle lobe with, deep, broad and paired Y-shaped impressions, their inner branches longer than the outer ones, terminating in deep pits (Fig. 8). Hind lobe robust, long and wide, ratio of maximum length to maximum width 0.38, with narrow median ridge. Venter of thorax not studied. Macropterous, forewing overlapping apex of abdomen, membrane lustrous, venation conspicuous, contrasting with membrane, not studied in detail. Fore legs long and slender. Ventral surface of fore-femur straight, dorsal surface slightly convex; ratio of length to maximum width 4.04. Fore tibia widening regularly towards apex, widest in a short apicitibial process, four times longer than wide. Bristle comb composed of approximately 32 setae. Apicitibial armature characteristic of Oncylocotis, formed by a compact group of seven spiniform setae; two short ventral setae, medial row of three long, strong setae, and two dorsal setae, slightly curved ventrally (Figs 11–12). Tarsal armature of two pairs (anterior and posterior) of spiniform setae, as illustrated in Fig. 11. Middle and hind legs very slender, without peculiarities.

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Figs 7–10. 7–8 – Oncylocotis stehliki sp.nov., male holotype; 9–10 – Oncylocotis lautereri sp.nov., female holotype; 7, 9 – head, dorsal view; 8, 10 – pronotum, dorsal view. Scale bar in mm.

Etymology. Patronymic, dedicated to our friend and colleague Dr. Jaroslav L. Stehlík for the occasion of his 90th birthday. Differential diagnosis. Oncylocotis stehliki sp.nov. differs from O. tasmanicus and O. saetosus in its smaller body size (5.3 mm; 6–7 mm in O. tasmanicus and O. saetosus); from O. lautereri sp.nov. in larger body size (4.25 mm in O. lautereri sp.nov.) and from all other described Australian species in its short, conspicuously curved vestiture of dorsum of body (longer, more erect in other species). Collecting circumstances. The single specimen of this species was attracted to light. Distribution. Australia, Northern Territory, Nitmiluk National Park.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 321 P. B AÒAØ & P. ŠTYS

Oncylocotis lautereri sp.nov. (Figs 2, 4, 6, 9–10, 13–14) Type material. Holotype: female, labelled: ‘AUSTRALIA, NT ; 63m alt. / Litchfield N.P. 20–27.11.2008 / TJAYNERA FALLS / 13°15′S, 130°44′E / L. Hovorka lgt.’ // ‘Holotype / Oncylocotis / lautereri sp. nov. / P. Baòaø & P. Štys det. 2013’ (MAGNT). Specimen is dry-mounted on rectangular card, right foreleg mounted separately on the same card and gold-coated for SEM photography. Diagnosis. Taking into account the newly-described Oncylocotis lautereri sp.nov. and all the described and undescribed species of Oncylocotis of which we are aware, O. lautereri sp.nov. may be recognized by the following unique combination of characters: legs very short and robust, ratio of fore-femur and fore-tibia length to their width less than 3, conspicuously longer setae on ventral surface of posterior lobe of head than those on the anterior, and very short antennae (ratio of antenna length to combined length of head and pronotum 0.7). Description. Submacropterous female (holotype). Measurements (in mm). Total body length – 4.24. Head (without neck). Total length – 0.81; posterior lobe, length – 0.25, posterior lobe, width – 0.46; distance of eye to apex of antennifer – 0.22; diatone (maximum width across eyes) – 0.42; dorsal synthlipsis (minimum interocular distance) – 0.28; eye, length – 0.17; gena, minimum width – 0.27; interocellar distance – 0.27; minimum distance ocellus–eye – 0.08. Labium. Total length – 0.68; segment I, length – 0.07; segment II, length – 0.09; segment III, length – 0.38; segment III, maximum height – 0.11; segment IV, length – 0.14. Antenna. Segment I, length – 0.14; segment II, length – 0.33; segment III, length – 0.29; segment IV, length – 0.38; segment II, basal width – 0.03, segment II, distal width – 0.07. Pronotum. Total length (maximum) – 0.84; collum, length (median) – 0.21, maximum width – 0.49; middle lobe, length (median) – 0.36, middle lobe, width (maximum) – 0.78; hind lobe, length (maximum) – 0.29, hind lobe, length (mediane) – 0.20, hind lobe, width (maximum) – 0.91. Fore leg. Femur, length – 0.71, femur, maximum width – 0.26; tibia, length – 0.56, tibia, maximum width – 0.20; tarsus, length – 0.13, tarsus, maximum width – 0.09; fore claws, length – 0.13. Fore wing. Maximum length – 2.11. Coloration. Body uniformly brown (Fig. 2), anterior lobe of head somewhat darker than posterior lobe. Legs and labium lighter. Abdomen dark brown. Sculpture. All body surfaces densely covered in very fine, minute granules, setigerous and non-setigerous. Vestiture. Dorsal and lateral surfaces of head with long, dense and “soft” semi-erect setae, slightly curved and directed forwards, except for the extreme rear part of the posterior lobe, where the setae are directed rearwards. Ventral surface of anterior lobe of head with erect setae pointing in various directions, but mostly directed rearwards, ventral face of posterior lobe with very long (in contrast to those on anterior lobe), erect setae, slightly directed forwards. Labium densely covered in semi-erect setae, mixed on all segments with longer setae. Antennae densely covered in semi-erect setae on all segments, on distal half of segment II and segments III and IV mixed with longer, more protruding setae. Thorax and fore wing veins densely covered with long, semi-erect to erect, slightly curved setae, directed rearwards. All surfaces of femora and tibiae covered

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Figs 11–14. Apicitibial armature of Oncylocotis spp. 11–12 – Oncylocotis stehliki sp.nov., male holotype; 13–14 – Oncylocotis lautereri sp.nov., female holotype; 13, 15 – apex of fore-tibia and fore-tarsus, anterior view; 14, 16 – apex of fore-tibia, lateroventral view.

in semi-erect, slightly curved setae of equal length; on dorsal and ventral surfaces and the posterior surfaces of the forelegs and anterior surfaces of mid- and hind legs mixed with long, erect, trichobothrium-like setae that stand out from the rest. Structure. Head (Figs 4, 6, 9) robust and wide, almost as long as pronotum. Eyes medium-sized, ocular index 4.0. Posterior lobe with broad, deep median impression, conspicuously transverse, almost twice as wide as long; ratio of length to width 0.54. Ocelli small, situated on small tubercles. Antennae very short, much shorter than head and pronotum together (ratio 0.7), antennal formula (longest segment first): IV-II-III-I. Labium (Fig. 4) short, reaching anterior margin of eye, labial formula (longest segment first): III-IV-II-I. Thorax. Pronotum (Fig. 10) narrow with shallow constrictions. Collum robust, wide, ratio of length to width 0.42, with well-developed precollum. Middle lobe long, robust, more than twice as wide as long (ratio 2.16); with broad, inversely T-shaped impression in the middle, reaching from collum-middle lobe constriction almost to posterior margin

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 323 P. B AÒAØ & P. ŠTYS of middle lobe. T-shaped impression with narrow median ridge continuing on pronotal hind lobe. Sublateral parts of middle lobe with pair of shallow, narrow, Y-shaped impressions. Hind lobe short, ratio of maximum length to maximum width 0.32, with narrow median ridge. Venter of thorax not studied. Submacropterous, forewing not reaching apex of abdomen, membrane lustrous, venation conspicuous, not studied in detail. Forelegs stout and robust. Ventral surface of fore-femur straight, dorsal surface conspicuously convex; ratio of length to maximum width 2.73. Fore-tibia widening regularly towards the apex, widest at a minute apicitibial process, where the ratio of length to maximum width is 2.8. Bristle comb composed of approximately 28 setae. Apicitibial armature very similar to O. stehliki sp.nov., formed by a group of seven spiniform setae; two short ventral setae (anterior one minute), medial row of three long, strong setae and two dorsal setae, slightly curved ventrally (Figs 13–14). Tarsal armature of two pairs (anterior and posterior) of spiniform setae, as illustrated in Fig. 13. Mid- and hind legs short and stout, without peculiarities. Etymology. Patronymic, dedicated to our colleague and friend Dr. Pavel Lauterer for the occasion of his 80th birthday. Differential diagnosis. Oncylocotis lautereri sp.nov. differs from all known Australian species of Oncylocotis in its smaller body size (4.25 mm, other species at least 5.0–5.5 mm or more) and in unusually short antennae (ratio of length to combined length of head and pronotum 0.7, in other species 0.9–1.0). Collecting circumstances. The single specimen of this species was attracted to light. Distribution. Australia, Northern Territory, Litchfield National Park.

Discussion Undescribed species of Oncylocotis abound in both the Australian and Papuan faunas. To describe all the taxa available is at present beyond us, particularly since some of the more broadly-distributed species exhibit surprisingly large and complex phenotypic variation within traits usually regarded as stable and diagnostic in the genus and family. This apparently intraspecific diversity seems to be a mix of variation – individual, sexual, population and cline. On the other hand, some less abundant species with restricted ranges appear to be easily diagnosed and two of them we do not hesitate to describe here, even though they are based on only single individuals.

Acknowledgement The authors would like to thank the two collectors, Stanislav Jákl and Ladislav Hovorka (both Prague) for loaning us these interesting specimens for our study. This paper appears through financial support provided to the Moravian Museum by the Ministry of Culture of the Czech Republic as part of its long-term conceptual development programme for research institutions (ref. MK000094862) and Grant of the

324 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Oncylocotis

Faculty of Science of Charles University in Prague (SVV-2013-267 201) (Petr Baòaø). Pavel Štys would like to extend his gratitude to Gerry Cassis (Sydney) for his hospitality while visiting the Australian Museum (Sydney) and for encouraging his interest in the Australian fauna.

References

CASSIS E. & GROSS G. F. 1995: Hemiptera: Heteroptera (Coleorrhyncha to ). In: HOUSTON W. W. K. & MAYNARD G. V. (eds.): Zoological Catalogue of Australia. Vol. 27.3A. CSIRO Australia, Melbourne, xv + 506 pp. JEANNEL R. 1942: Les Hénicocephalides. Monographie d’un groupe d’Hémiptères hématophages. Annales de la Société Entomologique de France 110: 273–368. STÅL C. 1856: Hemiptera från Kafferlandet. Öfversigt af Kongliga Svenska Vetenskaps-Akademiens Förhandlingar 12 (1855): 89–100. WESTWOOD J. O. 1837: Descriptions of several new genera of exotic hemipterous insects. Transactions of the Entomological society of London 2: 18–24.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 327–334, 2013

Two new species of the enigmatic genus Aquulavelia (Hemiptera: Heteroptera: ) from Nepal and Bhutan

HERBERT ZETTEL Department of Entomology, Natural History Museum, Burgring 7, 1010 Vienna, Austria; e-mail: [email protected]

ZETTEL H. 2013: Two new species of the enigmatic genus Aquulavelia (Hemiptera: Heteroptera: Veliidae) from Nepal and Bhutan. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 327–334. – Aquulavelia Thirumalai, 1999 is only known from the Indian subcontinent. Comparative notes on the genus and its type species, Aquulavelia occulta Thirumalai, 1999 from Arunachal Pradesh, north-eastern India, are given. Two new species are described: Aquulavelia stehliki sp.nov. from Nepal and A. lautereri sp.nov. from Bhutan. A key to species is provided. Keywords. , Veliidae, Microveliinae, true bugs, Aquulavelia, new species, key, Oriental Region, India, Nepal, Bhutan

Introduction Aquulavelia Thirumalai, 1999 is a genus of Microveliinae known only from the Indian subcontinent. The only taxonomic paper addressing Aquulavelia is the original description by THIRUMALAI (1999), who described the genus and its type species, Aquulavelia occulta, from Arunachal Pradesh, north-eastern India, and mentioned a second, undescribed species from Madhya Pradesh in central north India. At that time Dr. Thirumalai kindly donated a pair of paratypes to the collection of the Natural History Museum, Vienna. Although I have seen considerable Gerromorpha material from the southern parts of the Himalayas since, only two specimens of Aquulavelia have been identified, one male from Nepal and one female from Bhutan. Both specimens belong to a hitherto undescribed species and are here formally described. This study aims to draw the attention of hemipterologists and limnologists to this still-enigmatic genus.

Material and methods The Natural History Museum Vienna (NHMW) is the depository for all the type specimens studied. Inventory numbers are given. Label texts are cited. Terminology and mode of description follow earlier publications of the author. Measurements are in millimetres and refer to maximum width, median length (pronotum, tergites), or maximum length (other structures), excluding pilosity, measured in straight lines. Stacked digital images (Figs 1, 2) were taken with a Leica DFC camera attached to a Leica MZ16 binocular microscope and processed with the help of the Leica Application Suite. These were then stacked with ZereneStacker 64-bit and processed with Adobe Photoshop 7.0. For the production of line drawings a camera lucida attached to an

327 H. ZETTEL

Olympus BX40 compound microscope (Figs 3–8) and a Nikon SMZ800 binocular microscope (Figs 9, 10) was employed.

Taxonomy Aquulavelia Thirumalai, 1999 Aquulavelia Thirumalai, 1999: 205. Type species: Aquulavelia occulta Thirumalai, 1999, by original designation.

Diagnosis (modified from THIRUMALAI 1999). Species of Microveliinae with small and slender body (Figs 1, 2); body length 2.3–3.1 mm; only apterous morph known. Numerous long setae present on thoracic dorsum, abdominal tergites and laterotergites, antennomere 1, and femora. Head anteriorly deflected. Eye large, situated close to pronotum. Antenna long, filiform (Fig. 1); antennomere 4 longest, antennomere 2 shorter than antennomeres 1 and 3. Pronotum of apterous morph covering entire mesonotum and metanotum medially (Figs 1, 2). Lateral evaporatorium with short channel, ending close to metacetabula with a tuft of long setae. Legs simple; protibia of male with short grasping comb; claws long, falcate, inserting preapically; arolia bristle-like. Abdomen with subparallel sides in male, quite convex sides in female (Figs 1, 2). Some or all tergites with transverse rows of standing black setae at approximately distal third. Tergite 1 with conspicuously swollen sides and deep median impression. Sternites with more or less distinct, very narrow median impression (hardly visible in A. stehliki sp.nov.). Segment 8 of male inserted caudally, slightly depressed. Genitalia small. Parameres symmetrical, slender, falciform (Figs 7, 8). Proctiger in both sexes pointed at apex (Figs 3–6). Gonocoxa 1 of female plate-like, unmodified. Comparative notes and discussion. Although a phylogeny of Microveliinae of the world is not available, most genera – but not Microvelia Westwood, 1834 in the wider sense – seem to be justified by good sets of morphological characteristics. ANDERSEN & WEIR (2003) presented a table with 17 morphological characters for the 16 Australasian genera of Microveliinae (not including Aquulavelia) and some species groups of Microvelia; their cladogram of relationships indicates the polyphyly of Microvelia. THIRUMALAI (1999) compared Aquulavelia with Neoalardus Distant, 1912, Baptista Distant, 1903 and Lathriovelia Andersen 1989. Indeed, the overall similarities – namely the very elongated body – of Aquulavelia and Neoalardus are striking. However, Neoalardus differs significantly in the absence of a protibial grasping comb in the male and, in the apterous morph, by a medially visible metanotum, a totally differently structured tergite 1, and peculiar silverish spots on the sides of the metanotum and abdominal tergites 2, 4, and 6–8 (for Neoalardus see e.g. ZETTEL 1998). Males of Baptista usually have characteristic modifications of the profemur and the pregenital abdomen (see e.g. ANDERSEN 1989), except for the species of the Baptista collaris group from southeastern Asia which differ in a peculiar set of apomorphic characters, including protibial rows of stout, spine-like setae (in both sexes), sternite 7 of male medially angularly incised, and parameres distally dilated (ZETTEL 2004). The Malayan genus

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Lathriovelia, the macropterous species of which inhabit water-filled bamboo internodes, also have some extreme apomorphies, including small eyes distant from the pronotal margin and tergite 6 produced into paired lobes (KOVAC & YANG 2000). Last but not least, Aquulavelia should not be regarded as an aberrant clade of Microvelia, because it shares a caudal insertion of abdominal segment 8 of the male with most other genera of Microveliinae, while in typical species of Microvelia (and also in the species of Pseudovelia Hoberlandt, 1950) male segment 8 takes a more ventrocaudal position. Distribution. Nepal, Bhutan, India. Habitats. THIRUMALAI (1999) reported A. occulta from “dark pits of a slow flowing mountain stream”. Aquulavelia lautereri sp.nov. was collected on the gravel banks of a river around 30 metres wide, probably in puddles or in a small branch (M. A. Jäch, pers. comm.). Nothing is known about the collecting circumstances of A. stehliki sp.nov.

Key to species 1 Segment 1 of metatarsus longer than segment 2 (ca. 1.1–1.2×). Short appressed pilosity of all tergites relatively dense, surface hardly lustrous...... 2 – Segment 1 of metatarsus slightly shorter than segment 2 (0.95×). Short appressed pilosity on tergites 3–8 (of female) reduced, tergites lustrous. Bhutan...... A. lautereri sp.nov. 2 Antenna very long, antennomere 1 distinctly longer than head length (♂: 1.23×, ♀ 1.15×). Only tergites 1–3 and 7 of male with transverse rows of long setae...... A. occulta Thirumalai, 1999 – Antenna comparatively short, length of antennomere 1 subequal to head length (♂: 1.03×). All tergites of male with transverse rows of long setae...... A. stehliki sp.nov.

Aquulavelia occulta Thirumalai, 1999 (Figs 3, 5, 7, 9)

Aquulavelia occulta Thirumalai, 1999: 206 (original description); THIRUMALAI (2002): 72 (catalogue). Type material examined. Paratypes (1 apterous male, 1 apterous female): ‘INDIA: Arunachal Pradesh \ Upper Subasiri District \ road to Getha, Yingklong \ 700 m, 31 Aug. 1994 \ leg. G. Thirumalai’ (NHMW-Hemipt.- Inv.No. 000 014 428–429). Diagnostic characters. Body length of male c. 2.5–2.9, of female c. 2.8–3.1. Yellowish; dorsum of head, dorsum and sides of thorax (except a distinct yellow transverse mark near pronotal fore-margin), entire tergites 1 and 2, sides of following tergites, medial margins of laterotergites, and stripes on sides of sternites more or less infuscated. Antenna very long, in male 0.85×, in female 0.72× body length; antennomere 1 longer than head length (male: 1.23×, female: 1.15×); antennomere 2 slender and elongated, in male 0.7×, in female 0.6× as long as antennomere 1; also antennomeres 3 and 4 longer than in A. stehliki sp.nov. Profemur of male not incrassate; grasping comb of male 0.15×

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 329 H. ZETTEL as long as entire protibia (Fig. 9). Abdominal tergites entirely covered with relatively dense, appressed short pilosity, except in male narrow midlines of tergites 4–7 bold and lustrous (in the female the midlines are also slightly impressed, but pilose). In male tergites 1–3, and 7 with transverse rows of long, black, erect setae; in females all tergites with such setae. Proctigers of both sexes strongly pointed (Figs 3, 5). Paramere slender, moderately long, with slight bend near mid-length (Fig. 7). Comparative notes. See key and the notes for following species.

Aquulavelia stehliki sp.nov. (Figs 1, 4, 8, 10) Type material. Holotype (apterous male): ‘NEPAL: Harare Prov. \ Andhi Khola, S-63 \ 3.I.1994 \ leg. Suboth Sharma’ (NHMW-Hemipt.-Inv.No. 000 014 430). Description of apterous male. Measurements. Body length 2.35, width at metanotum 0.62. Head length 0.32, width 0.48. Distance between eyes 0.22. Lengths of antennomeres 1–4: 0.33, 0.21, 0.48, 0.55. Pronotum length 0.40, width 0.58. Lengths of leg segments: profemur 0.66, protibia 0.61, protarsus 0.24, mesofemur 0.77, mesotibia 0.73, mesotarsus 0.18+0.21, metafemur 0.91, metatibia 1.20, metatarsus 0.26+0.24. Abdomen width 0.54. Lengths of tergites 1–7: 0.17, 0.19, 0.19, 0.19, 0.20, 0.22, 0.21. Colour (Fig. 1). Trunk yellow to yellowish-brown. Head with large, dark mark between eyes. Pronotum dark brown, except a distinct yellow transverse mark near fore- margin; dark stripes on sides of tergites, medial margins of laterotergites, and sides of sternites only narrowly infuscated. Antennae and legs entirely pale yellow. Pilosity. Appressed pilosity covering the entire trunk; on some areas these hairs broader and silvery or golden lustrous, e.g. on entire pronotum and on tergites, but not forming clear marks. Long, black erect setae numerous; several such setae, of slightly varying length, on head dorsum, pronotum, and metanotum sides; on tergites forming not very regular transverse rows in posterior half, visible on tergites 1–7 (best seen in an anterodorsal aspect), 8, 6, 4, 4, 4, 3, 4 setae (as they are not symmetrically arranged, a few setae might be broken off); on all laterotergites a few standing setae of varying length; antennomere 1 with two long setae; profemur with several long setae in distal half of flexor side; mesofemur with a few setae at base of extensor side, and several setae more or less arranged in two rows on flexor side; metafemur with numerous setae more or less arranged in one row on extensor side and two rows on flexor side. Structure. Mostly as in A. occulta, with the following exceptions: profemur slightly incrassate in distal half; grasping comb 0.12× as long as entire protibia (Fig. 10). Laterotergites 2–6 convex, connexiva appearing swollen. Median impression on sternites very inconspicuous, only recognizable on sternites 5 and 6. Proctiger moderately pointed (Fig. 4). Paramere moderately long, evenly curved (Fig. 8). Comparative notes. Aquulavelia stehliki sp.nov. differs from A. occulta in its clearly shorter antennae. The entire antenna length is about 0.67× body length in the male of A. stehliki sp.nov., whereas it reaches 0.85× body length in the male of A. occulta (only

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Fig 1. Aquulavelia stehliki sp.nov., holotype, male (abdominal segment 8 and genitalia dissected).

0.72× in the female). The male of A. occulta has a reduced number of long, erect setae on the tergites, especially tergites 4–6 have 0–1 erect setae. In the male of A. stehliki sp.nov., complete transverse rows of 3–8 standing setae are present. In the male of A. occulta the narrow midlines of tergites 4–7 are bold and lustrous, whereas in A. stehliki sp.nov. all tergites are entirely pilose. Moreover, A. stehliki sp.nov. is slightly smaller than A. occulta, its profemur slightly incrassate, and its protibial grasping comb slightly shorter. The genitalia of the two species are distinctive: the paramere of A. stehliki sp.nov. is more regularly curved and its proctiger is less pointed. Etymology. This species is dedicated to Dr. Jaroslav Stehlík for the occasion of his 90th birthday.

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Fig 2. Aquulavelia lautereri sp.nov., holotype, female.

Aquulavelia lautereri sp.nov. (Figs 2, 6) Type material. Holotype (apterous female): ‘BHUTAN: Sarpang Prov. \ 11 km NW Sarpang \ Bhur Khola, ca. 350 m \ 26°55′23″N 90°23′51″E \ 27.11.2005, leg. M. Jäch (30)’ (NHMW-Hemipt.-Inv.No. 000 014 431). Description of apterous female. Measurements. Body length 2.69, width at metanotum 0.74. Head length 0.37, width 0.52. Distance between eyes 0.26. Lengths of antennomeres 1–2 (3 and 4 missing): 0.37, 0.26. Pronotum length 0.43, width 0.62. Lengths of leg segments: profemur 0.74, protibia 0.64, protarsus 0.29, mesofemur 0.86, mesotibia 0.84, mesotarsus 0.22+0.27, metafemur 1.05, metatibia 1.32, metatarsus 0.27+0.29. Abdomen width 0.76. Lengths of tergites 1–7: 0.20, 0.22, 0.23, 0.23, 0.26, 0.27, 0.30.

332 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Aquulavelia

Figs 3–10. Diagnostic structures of Aquulavelia spp. 3–6 – proctigers: 3 – A. occulta Thirumalai, 1999, male; 4 – A. stehliki sp.nov., male; 5 – A. occulta, female; 6 – A. lautereri sp.nov., female. 7–8 – parameres: 7 – A. occulta; 8 – A. stehliki sp.nov. 9–10 – forelegs of males: 9 – A. occulta; 10 – A. stehliki sp.nov. Pilosity omitted in Figs 3–6, partly omitted in Figs 9 and 10.

Colour (Fig. 2). Ground colour of trunk yellow. Blackish-brown: dorsum of head and thorax, except a distinct yellow transverse mark near pronotal fore-margin, metapleura, abdominal tergites except for round yellow marks on tergites 1–6 and a narrow yellow midline on tergite 7, medial halves of all laterotergites, and broad continuous stripes on sides of sternites. Antennae and legs entirely brownish-yellow, bases of legs pale. Pilosity. Appressed pilosity present on entire trunk, but sparse on tergites 3–8 and lateral parts of laterotergites and sternites; these areas therefore lustrous; on some areas, these hairs broader and silvery-lustrous, i.e., on head between eyes, entire pronotum, sides of tergites and medial margins of laterotergites, but not creating clear markings. Long, black erect setae numerous; several such setae, of slightly varying length, on head dorsum, pronotum, and metanotum sides; on some tergites forming slightly irregular transverse rows in posterior half, visible on tergites 1–8 (best seen in anterodorsal aspect), 14, 6, 5, 2, 0, 0, 11, 8 setae (as they are not symmetrically arranged, a few setae might be broken off); on all laterotergites a few setae of varying length; antennomere 1 with two long setae; profemur with two rows of long setae on flexor side; mesofemur and

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 333 H. ZETTEL metafemur each with numerous long setae, more or less arranged in several rows, but absent from anterior faces of femora. Structure. Mostly as in A. occulta, except medial impression of tergite 1 shallower and apical spine of proctiger slightly shorter (Fig. 6). Comparative notes. Aquulavelia lautereri sp.nov. differs from A. occulta in segment 1 of the metatarsus slightly shorter than segment 2 (longer in A. occulta and A. stehliki sp.nov.) and in antennal segment 1 as long as head length (longer in A. occulta). In the female (male unknown) tergites 3–8 are lustrous and their appressed pilosity is reduced; the proctiger spine is shorter (comp. Figs 5 and 6). Black coloration is more developed on the abdomen of A. lautereri sp.nov. than in A. occulta. Whereas in A. lautereri sp.nov. clearly delimited black stripes are developed at lateral parts of tergites, medial halves of laterotergites, and sublaterally on sternites, only weak infuscations are present in the same areas in A. occulta. Etymology. This species is dedicated to Dr. Pavel Lauterer for the occasion of his 80th birthday.

Acknowledgements I am very grateful to Dr. Manfred A. Jäch (Natural History Museum, Vienna), Prof. Dr. Suboth Sharma (Kathmandu University), and the late Dr. G. Thirumalai (Zoological Survey of India, Chennai) for providing specimens for this study. I also thank Tony Long (Svinošice) for helping to work up the English.

References

ANDERSEN N. M. 1989: The Old World Microveliinae (Hemiptera: Veliidae). II. Three new species of Baptista Distant and a new genus from the Oriental region. Entomologica Scandinavica 19: 363–380. ANDERSEN N. M. & WEIR T. A. 2003: The genus Microvelia Westwood in Australia (Hemiptera: Heteroptera: Veliidae). Invertebrate Systematics 17: 261–348. KOVAC D. & YANG C. M. 2000: Revision of the Oriental bamboo-inhabiting semiaquatic bug genus Lathriovelia Andersen, 1989 (Heteroptera: Veliidae) with description of L. rickmersi, new species, and notes on the genus Baptista Distant, 1903. Raffles Bulletin of Zoology 48(1): 153–165. THIRUMALAI G. 1999: A new genus of Microveliinae (Hemiptera: Heteroptera: Veliidae) from north eastern India with a checklist of the Indian species of the subfamily. Records of the Zoological Survey of India 97(3): 205–211. THIRUMALAI G. 2002: A check list of Gerromorpha (Hemiptera) from India. Records of the Zoological Survey of India 100(1–2): 55–97. ZETTEL H. 1998: On the genus Neoalardus Distant 1912 (Heteroptera: Veliidae). Linzer Biologische Beiträge 30(2): 595–599. ZETTEL H. 2004: Baptista hoedli sp.nov. (Insecta: Heteroptera: Veliidae) from Laos, and definition of the B. collaris species group. Annalen des Naturhistorischen Museums in Wien, Serie B 105 (2003): 441–445.

334 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 335–346, 2013

A new species and new synonymy in Limnocoris (Hemiptera: Heteroptera: ) from Brazil

NICO NIESER1, PING-PING CHEN1 & ALAN LANE DE MELO2 1 Department of Terrestrial Zoology, Naturalis Biodiversity Centre, Postbus 9517, 2300 RA Leiden, The Netherlands; e-mail: [email protected], [email protected] 2 Departamento de Parasitologia, Instituto de Ciencias Biologicas, Universidade Federal de Minas Gerais, Av. Antonio Carlos 6627, 31270/901 Belo Horizonte, Minas Gerais, Brasil; e-mail: [email protected]

NIESER N., CHEN P.-P. & DE MELO A. L. 2013: A new species and new synonymy in Limnocoris (Hemiptera: Heteroptera: Naucoridae) from Brazil. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 335–346. – Limnocoris lautereri sp.nov. is described from Minas Gerais, Brazil. A lectotype is designated for Limnocoris volxemi (Lethierry, 1877), and L. maculiceps Montandon, 1897, syn.nov., is established as a junior subjective synonym of L. volxemi. Keywords. Heteroptera, Naucoridae, Limnocoris, aquatic bugs, lectotype designation, new synonymy, new species, taxonomy, Neotropical Region, Brazil

Introduction The genus Limnocoris Stål, 1860 is currently placed in the monotypical subfamily Limnocorinae Stål, 1876, under the family Naucoridae Leach, 1815. The genus comprises 71 known species, including the new one described here, of which 30 occur in Brazil, 13 in Minas Gerais, and 6 in Amazonas (NIESER & LOPEZ RUF 2001, MOREIRA et al. 2011, this work). Limnocoris is essentially a Neotropical genus, although one somewhat aberrant species (L. moapensis La Rivers, 1950) lives in certain warm springs in Nevada, U.S.A., and another species’ distribution reaches Texas (HENRY & FROESCHNER 1988). The greatest diversity of species is found in the Andes mountain range (NIESER et al. 1993) and in the area from south-eastern Brazil to northern Argentina (NIESER & LOPEZ RUF 2001). In Minas Gerais, most limnocorids live in small to medium-sized streams with slow to moderate current and a bottom of fine to coarse sand or small pebbles. The bugs appear to be nocturnal, hiding in the sandy substratum by daytimeh; however, nothing definite is known of their way of life. The most commonly found species in the state, L. volxemi Lethierry, 1877, has usually been caught among vegetation at the virtually still edges of streams, and occasionally in still-water pools in marshes as well. The first revision of the genus was undertaken by MONTANDON (1898), who recognized 17 species. DE CARLO (1951) published photographs of the 13 species present in the collection of the Museo Argentina de Ciencias Naturales, mostly from Argentina and southern Brazil. Subsequently, J. A. de Carlo and I. La Rivers described a number of

335 N. NIESER ET AL. new species, which were summarized in papers by LA RIVERS (1971, 1974, 1976). NIESER et al. (1993) described five new species from Departamento Valle del Cauca, Colombia, and later NIESER & LOPEZ RUF (2001) revised the genus for the southern area of South America, adding 10 new species, pointing out some synonymies, and presenting a complete checklist of species. In the course of work on Heteroptera samples collected from Amazonas in Brazil, a previously unknown species of Limnocoris was disclosed, which is described herein. In addition, the correct taxonomic status of L. volxemi is studied and confirmed. We dedicate this work to the outstanding hemipterists RNDr. Pavel Lauterer and RNDr. Jaroslav L. Stehlík; also to express gratitude for the invaluable help to Ping-Ping Chen in her study on Homoptera.

Material and methods The material examined is deposited in the following collections: DPIC ...... Departamento de Parasitologia, Instituto de Ciencias Biologicas, Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brasil ISNB ...... Département d’Entomologie, Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium MNHN ...... Muséum national d’Histoire naturelle, Paris, France NCTN ...... N. Nieser & P.-P. Chen collection, Tiel, The Netherlands

In our approach to the study and to terminology, we follow NIESER & LOPEZ RUF (2001). The term “synthlipsis” is the smallest distance between the eyes at the rear. The detailed positions at which measurements were taken appears in Figs 24–25; all measurements are given in mm and presented as the mean for certain more important measurements, followed by the range in parentheses. Means are based on five specimens for each sex. The sternal carinae are described with the ventral side upward. The black-and-white line illustrations were made with the aid of a camera lucida on a Leitz stereo microscope, and an Olympus monocular microscope. Habitus photos were taken with a Nikon D700 body with AF 60 mm f2.8D Micro-Nikkor lens, and a Leica 500, and worked up with Adobe Photoshop CS2, version 9.0.

Results Limnocoris lautereri sp.nov. (Figs 3, 4, 5, 10, 13, 16, 17)

Type material. Holotype: ♂ (brachypterous), Brasil: Amazonas: Pitinga, 00°47′28.7″S, 60°04′12.2″W, Estaude de Tiros, abaixo da corredeira, 2.iv.2000, leg. D. L. V. Pereira (DPIC). PARATYPES: 14 ♂♂ 14 ♀♀ (all brachypterous), same data as holotype (7 ♂♂ 7 ♀♀ DPIC, 7 ♂♂ 7 ♀♀ NCTN). Description (based on specimens stored in 70% ethanol). Generally a small, light brown, broadly oval species (Fig. 3). Dimensions. Body length: ♂♂ 6.6 (6.5–6.8) {HT 6.6}, ♀♀ 6.6 (6.5–6.8), width across embolia: ♂♂ 4.6 (4.5–4.8) {HT 4.8}, ♀♀ 4.7 (4.6–4.8), width across connexiva: ♂♂ 4.2 (4.1–4.4) {HT 4.4}, ♀♀ 4.3 (4.2–4.4), anterior width of vertex: ♂♂ 1.32

336 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and synonymy in Limnocoris

Figs 1–3. 1–2 – Limnocoris volxemi (Lethierry, 1877): 1 – lectotype, male; body length 9.0 mm; 2 – paralectotype, female; body length 9.5 mm. 3 – Limnocoris lautereri sp.nov., paratype, male; body length 6.2 mm.

(1.30–1.36) {HT 1.34}, ♀♀ 1.35 (1.30–1.40), width of pronotum ♂♂ 4.04 (3.90–4.16) {HT 4.16}, ♀♀ 4.06 (3.98–4.17), median length of pronotum: ♂♂ 1.31 (1.24–1.35) {HT 1.35}, ♀♀ 1.36 (1.30–1.40), lateral length of pronotum: ♂♂ 1.65 (1.56–1.72) {HT 1.64}, ♀♀ 1.66 (1.62–1.72), median length of head + pronotum: ♂♂ 2.03 (1.88–2.18) {HT 2.18}, ♀♀ 2.10 (1.97–2.15). Colour. Dorsally brown, lateral parts of pronotum and embolia with an orange hue. Head orange with variable, indistinct darker markings medially, eyes black. Scutellum darker than clavus and corium, right membrane with dark vermiculate pattern. Legs and prosternum orange-brown, meso- and metasternum velvety grey, metasternum with an orange tinge towards the rear. Abdominal venter brown to grey, laterally lighter. Structure. Anterior margin of head following the curvature of pronotum and eyes, not projecting. Median mark of interoculus indistinct, anterior width of vertex 1.3 times synthlipsis (1.34 / 1.04); head shorter than medial length of pronotum (1.17 / 1.34); length of eye 1.9 times its width (0.91 / 0.49); anteclypeus projecting 0.1 forward beyond labrum (Fig. 30). Labrum parallel-sided in basal half and converging to an acute tip in apical half, densely covered in dirty-white, short, adpressed, somewhat scale-like hairs; its basal width is equal to its median length (0.42 / 0.42). Pronotum with lateral angles obtusely rounded, maximum width 0.15–0.30 removed from posterior margin, central area lumpy but not distinctly elevated, lateral margin finely serrate, barely visible to

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 337 N. NIESER ET AL.

Figs 4–9. Limnocoris spp., apex of abdomen, scale 1 mm. 4, 6, 8 – male, in dorsal view: 4 – L. lautereri sp.nov., paratype; 6 – L. pusillus Montandon, 1897; 8 – L. illiesi De Carlo, 1967. 5, 7, 9 – female, in ventral view: 5 – L. lautereri sp.nov., paratype, segment VIII removed; 7 – L. pusillus; 9 – L. illiesi.

338 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and synonymy in Limnocoris

Figs 10–15. Limnocoris spp., mesosternum. 10–12 – Mesosternal carina, lateral view from the right side, scale 1 mm: 10 – L. lautereri sp.nov., paratype; 11 – L. pusillus Montandon, 1897; 12 – L. illiesi De Carlo, 1967. 13–15a – mesocarinal fossa in ventral view: 13 – L. lautereri sp.nov., paratype; 14 – L. illiesi De Carlo, 1967; 15 – L. pusillus Montandon, 1897; 15a – L. volxemi (Lethierry, 1877).

indiscernible in dorsal view, about 30 teeth per mm in ventral view; maximum width less than twice anterior width (4.05 / 2.25), and 2.4 times the length of lateral section (4.05 / 1.66). Scutellum 2.3 times as wide as its median length (2.20 / 0.97) and 1.6 times as long as the claval commissure (0.97 / 0.62). Hemelytra leaving a narrow (0.10–0.15 wide) lateral strip of abdomen exposed. Embolium (Fig. 3) with lateral margin only slightly convex, smoothly curved, posterolateral angle virtually absent, its lateral margin indistinctly serrate, in most specimens hardly visible in dorsal view, about 26 teeth per mm in ventral view, width across embolium distinctly larger than maximum width across connexiva (4.7 / 4.3), claval suture absent, right membrane coriaceous, broadly rounded, covering left membrane, membranal suture absent. Left membrane well-developed, membranous. Hind wings strongly reduced, whitish, membranous, reaching over the base of abdomen. Covered part of abdomen waxy medium-rufous brown. Connexival segments not indented, forming a continuous lateral margin to abdomen, last segment with a small spine at the rear. Lateral margin of connexiva distinctly serrate, on segment IV with 34–37 teeth per mm. Prosternal carina with a small knob at the front, slanting downwards gradually towards the rear. Mesosternal carina (Figs 10, 13) well developed, with a low, blunt tooth at the front and a larger tooth at the central part sloping upwards and towards the rear, which has a very shallow, oval fossa; rear of central part sloping downwards abruptly. Metasternal carina with a rhomboid central part with a very shallow

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 339 N. NIESER ET AL.

Figs 16–23. Limnocoris spp., pseudoparameres. 16–17 – L. lautereri sp.nov., paratype, scale 0.1 mm (16 – right, in dorsal view; 17– left, in dorsal view). 18–20 – L. pusillus Montandon, 1897, scale 0.25 mm (18 – left, in dorsal view; 19 – right, inner face; 20 – right, in dorsal view). 21–23 – Limnocoris illiesi De Carlo, 1967, scale 0.25 mm (21 – left, in dorsal view; 22 – right, inner face; 23 – right, in dorsal view).

fossa, strongly slanting downwards at both front and rear. Fore-femur twice as long as wide (♂♂ 1.60 / 0.80, ♀♀ 1.55 / 0.74). Middle femur 5.4 times as long as wide (1.62 / 0.30), and 1.4 times as long as middle tibia (1.62 / 1.16). Hind femur 6.5 times as long as wide (2.22 / 0.34), and about as long as hind tibia (2.22 / 2.12). Male. Genital segments as in Fig. 4, proximal projection of right lobe of tergite VI slight. Pseudoparameres as in Figs 16 and 17. Female. Genital operculum (Fig. 5) with broadly rounded subapical projections giving the basal part of its lateral margin a concave appearance. Comparative notes. In its small size, this species is similar to L. pusillus Montandon, 1897 and L. illiesi De Carlo, 1967. Both of these species are different in mesocarinal shape (Figs 11, 12), and have very narrow mesocarinal fossae (Figs 14, 15), whereas L. lautereri sp.nov. has a different shape of mesocarina (Fig. 10) and a wide, shallow mesocarinal fossa (Fig. 13). The genital opercula of the females of both L. pusillus and L. illiesi (Figs 7, 9) are more evenly convergent at the rear than those in females of L.

340 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and synonymy in Limnocoris

Figs 24–29. Limnocoris volxemi (Lethierry, 1877), scale 1 mm, unless otherwise indicated: 24 – outline of body showing the measurements and structures employed herein; 25 – right foreleg in dorsal view; 26 – right hind tibia in dorsal view; 27 – apex of abdomen in dorsal view showing the expanded segments and genital capsule; 28 – apical part of aedeagus, scale 0.5 mm; 29 – female apex of abdominal tip, showing the operculum. Abbreviations: a = aedeagus; cc = claval comissure; le = length of eye; l = length; ll = length of lateral section of pronotum; p = paramere; pp = proximal projection; ps = pseudoparamere; r = stridulatory ridge, scale 1 mm; s = synthlipsis; v = anterior width of vertex; w = width; we = width of eye.

lautereri sp.nov. (Fig. 5); in addition, the operculum of the female of L. illiesi is obtusely rounded caudally (Fig. 9). The right pseudoparameres (Figs 16, 19–20, 22–23) and the left pseudoparameres (Figs 17, 18, 21) of L. lautereri sp.nov., L. pusillus and L. illiesi are also different. Etymology. This species is named in honour of Dr. Pavel Lauterer, the famous hemipterist from the Czech Republic, for his excellent contributions to the knowledge of Hemiptera, especially to the taxa of Psylloidea and Auchenorrhyncha. Distribution. So far known by only the type series from Brazil (Amazonas).

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Limnocoris volxemi (Lethierry, 1877) (Figs 1, 2, 15a, 24–29) Borbocoris volxemi Lethierry, 1877: 41. Limnocoris maculiceps Montandon, 1898: 417, 424–425 (syn.nov.) Limnocoris maculiceps: DE CARLO (1951: 44–45); NIESER & MELO (1997: 55); NIESER & LOPEZ RUF (2001: 281–283).

Type material examined. Limnocoris volxemi: Lectotype (here designated), ♂ (Fig. 1), and paralectotype, ♀ (Fig. 2) (here designated), both in brachypterous forms. Very little information was given by the original labels as: “PORTUGAL”: coll. Camille van Volxem (ISNB). Limnocoris maculiceps: Lectotype, brachypterous female, bearing labels: Museum Paris, Matto Grosso, de Castelneau 13-47 (printed) and Limnocoris maculiceps Montandon, type 1897 (handwritten, apparently by Montandon) and Lectotype dsg. Nieser & Lopez Ruf (printed on red paper). Paralectotype, macropterous male, bearing labels: Museum Paris, Matto Grosso, de Castelnau 13-47 (printed), and Limnocoris maculiceps cotype 1897 Montandon (handwritten), and Paralectotype (printed on blue paper) (MNHN, designated by NIESER & LOPEZ RUF 2001). Additional material examined. Brasil: Minas Gerais: Serra do Cipó, km 122, large and virtually still pothole in stream, surrounded by marsh, 24.ii.1996, leg. N. Nieser, N9603, 3 ♂♂ 3 ♀♀; Perdizes, small stream just south of village, 19.iii.1996, leg. N. Nieser & A. Pelli N9630, 3 ♂♂ 3 ♀♀; São Roque de Minas, Rio Peixe at camping site, 7.xi.1997, leg. N. Nieser & A. Pelli, N9724, 3 ♂♂ 3 ♀♀, all brachypterous (all in NCTN). Redescription. Brachypterous form (based on freshly-pinned specimens from the Serra da Canastra and surroundings). Generally a medium-sized, broadly oval medium- to dark brown species. Dimensions. Body length in male 8.60 (8.30–9.00), in female 8.80 (8.30–9.40), width of body in male 6.36 (6.29–6.50), in female 6.37 (6.10–6.60); anterior width of vertex in male 1.84 (1.80–1.90), in female 1.86 (1.78–1.98); width of pronotum in male 5.35 (5.20–5.50), in female 5.38 (5.15–5.65), median length of pronotum in male 1.89 (1.75–1.98), in female 1.90 (1.80–2.00); lateral length of pronotum in male 2.43 (2.38–2.50), in female 2.47 (2.42–2.50); median length of head + pronotum in male 3.18 (3.10–3.28), in female 3.24 (3.20–3.30). Colour. Light to medium-brown with variable darker mottling, which may become confluent resulting in darker specimens; with light patches laterally on interoculus, submedially and laterally on pronotum and laterally on scutellum, interoculus with a characteristic dark mark, broad towards the front, narrowing towards the rear (Figs 1, 2); eyes dark brownish-grey. Embolium in lighter specimens usually unmarked, yellowish; in darker specimens greyish, contrasting only a little with corium. Membrane concolorous and somewhat more densely mottled than corium. Exposed part of connexival segments in dorsal view yellowish with a distinct dark patch in anterior half. Dorsum of abdomen usually velvety yellowish to orange, occasionally blackish. Venter usually yellowish to light orange-brown in darkest specimens, a pair of dark patches anterolaterally on propleurae, embolium infuscated and connexival segments with dark patches anteriorly, which are larger in some specimens, forming an ill-defined infuscate lateral band, occasionally venter dark grey. Legs yellowish. Structure. Anterior margin of vertex following the curvature of pronotum and eyes, not protruding. Head distinctly declining anteriorly, anterior width of vertex 1.3 times the synthlipsis (1.85 : 1.44), head distinctly shorter than median length of pronotum (1.53 : 1.89), length of one eye slightly less than twice its width (1.24 : 0.65). Anteclypeus

342 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and synonymy in Limnocoris

Fig. 30. Limnocoris lautereri sp.nov., paratype (scale 1 mm): head, in lateral view, showing the anterior margin of head with anteclypeus projecting beyond labrum.

projecting 0.15–0.20 forward of labrum. Labrum parallel-sided in basal part, apical half triangular with a comparatively sharp tip, basal width more than median length (0.60 : 0.55). Pronotum posterolaterally rounded, angles just anterior of a line through the central part of the posterior margin (0.15–0.20), maximum width of pronotum distinctly less than twice its anterior width (5.37 : 3.40) and 2.2 times its length (5.37 : 2.45), lateral margin nearly straight, in posterior half weakly serrate ventrally with about 30 teeth per mm. Granulations on dorsal surface of body fine, lying completely on top. Scutellum clearly over twice as wide at base than its median length (3.02 : 1.29), and much less than twice as long as claval commissure (1.29 : 0.84). Hemielytra leaving only a narrow strip (0.2) of connexiva exposed laterally. Embolium slightly convex dorsally, lateral margin nearly straight but distinctly diverging towards the rear in anterior half, smoothly sinuate at the rear, lateral margin serrate, with approximately 20 teeth per mm; maximum width across connexiva 6.37 : 5.90. Claval suture indicated, right membranal suture absent, membrane coriaceous, broadly rounded, covering most of left membrane. Part of left membrane covered by right membrane, membranous, dark velvety-brown to blackish, broadly rounded, membranal suture absent; the lateral exposed part of left membrane 0.5 times or less wide at base, coriaceous and concolorous with corium. Hind wings reduced, reaching halfway along the third abdominal tergite, transparent to whitish-opaque. Connexival segments not indented, forming a smooth outer edge to abdomen, posterolateral angle of last connexival segment with a short spine (Fig. 27). Connexiva l laterally strongly serrate, connexiva 4 with about 15 teeth per mm. Prosternal carina of average elevation,

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 343 N. NIESER ET AL. distinctly sloping towards the rear, its bifid anterior tip indistinct in several specimens. Mesosternal carina with central part elevated, with a distinct fossa usually covered in hairs (Fig. 15a), its posterior part without teeth, short and steep. Metasternal carina poorly developed, a barely-elevated tubercle centrally on metaxiphus. Fore-femur (Fig. 25) 1.9 times as long as wide (2.18 : 1.16). Middle femur 5.2 times as long as wide (2.22 : 0.40) and 1.4 times longer than middle tibia (2.22 : 1.57). Hind femur 6 times as long as wide (3.07 : 0.49) and about as long as hind tibia (Fig. 26) (3.07 : 2.96). Male genital segments as in Figs 27–28. The right lobe of tergite 6 with a distinct projection proximally (Fig. 27). Pseudoparameres (Fig. 27) short and stout. Aedeagus (Fig. 28) with the apical widened part partly membranous. Female operculum as in Fig. 29. Macropterous form (based on NIESER & LOPEZ RUF 2001). Very similar to brachypetrous form except for hind wings fully developed. Light brownish opaque. Hemielytra with claval suture and nodal furrow well developed, right membrane only slightly larger compared with brachypterous form; left membrane and scutellum distinctly more developed; basal width of scutellum 3.20, median length 1.53; length of claval commisure 0.67; body length in male 7.8–8.7, in female 8.9, width in male 5.2–6.3, in female 6.6, median length of head + pronotum in male 3.40, in female 3.30. Fifth instar nymph. Dimensions. Length of body 8.20, width 6.30, anterior width of vertex 1.43, width of pronotum 5.40, median length of pronotum 1.17, length of lateral section of pronotum 2.05, median length of head + pronotum 2.45. Colour: Yellowish with variable, usually extensive brown patches formed by confluence of small dots. In specimens with well-developed brown colour pattern, the patch on the interoculus is as in adults, pronotum with a median and two lateral longitudinal bands on disk and a large posterior patch laterally at level of posterior half of eyes. Mesoalinotum irregularly patched, patches nearly confluent centrally. Metanotum and dorsum of abdomen with a tendency to become uniformly brown with distinct yellow patches laterally, forming quite a regular alternating light and dark pattern laterally on connexiva. Venter medially brown, laterally yellow with the dorsal pattern translucent. Structure. Lateral margin of connexiva distinctly serrate, finer than in the adult with about 22 serrations/mm on connexiva IV. Distribution. To date, it appears endemic to Brazil (Minas Gerais, Mato Grosso, Paraná, Santa Catarina). This is the most commonly found species in Minas Gerais, collected in 15 municipalities. Discussion. Borbocoris volxemi was described by LETHIERRY (1877) on the basis of two males and two females from Portugal, collected by C. van Volxem. Borbocoris is a subsequent incorrect spelling of Borborocoris Stål, 1861, and the latter name was synonymized with Limnocoris Stål, 1860 by MONTANDON (1898). Van Volxem collected the water bugs both in Portugal and in Brazil, and some of his Brazilian specimens appear to have become mixed with his Portuguese samples. Through the kindness of J. Constant and P. Grootaert (Royal Belgian Institute for Natural Sciences, Museum, Department of Entomology at Brussels, Belgium), it was possible to study the male and female syntypes of B. volxemi. These specimens had been

344 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species and synonymy in Limnocoris studied previously by G. Zimmermann, who was at that time working at the University of Marburg, Germany. The specimens bear red labels: “Lectotype / Paralectotype désigné par G. Zimmermann 1984”. Both specimens are glued on cards, the lectotype (Fig.1) is a male with dissected genitalia, and the paralectotype is a female (Fig. 2). Both specimens bear additional labels: “Portugal coll. Camille Van Volxem / Borborocoris volxemi Lethierry / syntype (red lettering)”. G. Zimmermann (in litt.) wrote that he had neither published this lectotype designation nor did he plan to do so. Therefore, the lectotype designation is validated here, and a note has been added under the lectotype labels “validated by Nieser, Chen et de Melo, 2013”. Considering the size, colour pattern, shape, male genitalia, female genital operculum, and mesosternal carina, L. maculiceps specimens examined are identical with those of the lectotype and paralectotype of L. volxemi, so we here synonymize L. maculiceps Montandon, syn.nov., with L. volxemi (Lethierry).

Acknowledgements We are indebted to Jérôme Constant and Patrick Grootaert (Royal Belgian Institute for Natural Sciences, Museum, Brussels, Belgium) for managing the loan of material for our study; and to Gerhard Zimmerman (Wuppertal, Germany) for the discussion on the species status of Limnocoris volxemi (Lethierry, 1877). We sincerely thank Thor-Seng Liew (Naturalis, Leiden, The Netherlands) for helping with the photography of the bugs. Finally, for proof reading and critical review, both of which have improved the content of this paper, we sincerely thank Petr Kment (Prague, Czech Republic) and Felipe Moreira (Rio de Janeiro, Brazil).

References

DE CARLO J. A. 1951: Género Limnocoris Stal (Hem. Naucor.). Misiones de Estudios de Patologia Regional Argentina 22: 41–51. HENRY T. J. & FROESCHNER R. C. (eds.) 1988: Catalog of the Heteroptera or true bugs of Canada and the continental United States. Brill, Leiden, New York, xix + 958 pp. LA RIVERS I. 1971: Catalogue of taxa described in the family Naucoridae (Hemiptera). Memoirs of the Biological Society of Nevada 2: 65–99. LA RIVERS I. 1974: Catalogue of taxa described in the family Naucoridae (Hemiptera). Supplement 1. Occasional Papers of the Biological Society of Nevada 38: 1–17. LA RIVERS I. 1976: Catalogue of taxa described in the family Naucoridae (Hemiptera). Supplement 2. Occasional Papers of the Biological Society of Nevada 41: 1–17. LETHIERRY L. 1877: Relevé des Hémiptères recueillis en Portugal et en Espagne par M. C. van Volxem. Annales de la Société Entomologique de Belgique 20: 34–43. MONTANDON A. L. 1898: Hemiptera cryptocerata. Fam. Naucoridae. – Sous-fam. Limnocorinae. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 48: 414–425. MOREIRA F. F. F., BARBOSA J. F., RIBEIRO J. R. I. & ALECRIM V. P. 2011: Checklist and distribution of semiaquatic and aquatic Heteroptera (Gerromorpha and ) occurring in Brazil. Zootaxa 2958: 1–74. NIESER N., GONZÁLEZ R. & EICHELKRAUT K. 1993: Nuevas especies de Naucoridae Fallen, (Heteroptera: Nepomorpha). Boletín del Museo de Entomología de la Universidad del Valle 1: 1–11.

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NIESER N. & LOPEZ-RUF M. 2001: A review of Limnocoris Stål (Heteroptera: Naucoridae) in southern South America East of the Andes. Tijdschrift voor Entomologie 144: 261–328. NIESER N. & MELO A. L. DE 1997: Os Heterópteros Aquáticos de Minas Gerais. Ed. UFMG, Belo Horizonte, Brasil, 180 pp. STÅL C. 1860: Bidrag till Rio Janeiro-Traktens Hemipter-Fauna. Kungliga Svenska Vetenskapsakademiens Handlingar 2(7) (1858): 1–84. STÅL C. 1861: Nova methodus familias quasdam Hemipterorum disponendi. Öfversigt af Kungliga Svenska Vetenskapsakademiens Förhandlingar 18: 195–212.

346 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 347–361, 2013

Two new species of thread-legged assassin bugs from Australia (Hemiptera: Heteroptera: Reduviidae: )

DÁVID RÉDEI Institute of Entomology, Faculty of Life Sciences, Nankai University, Weijin Road 94, 300071 Tianjin, China & Department of Zoology, Hungarian Natural History Museum, H-1088 Budapest, Baross u. 13, Hungary; e-mail: [email protected]

RÉDEI D. 2013: Two new species of thread-legged assassin bugs from Australia (Hemiptera: Heteroptera: Reduviidae: Emesinae). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 347–361. – Two new species of thread-legged assassin bugs (Hemiptera: Heteroptera: Reduviidae: Emesinae) are described from Australia: stehliki sp.nov. (Leistarchini) from Western Australia: Derby and Bargylia lautereri sp.nov. () from Queensland: Cooktown. The type species of the genus Bargylia Stål, 1866 is discussed. Keywords. Heteroptera, Reduviidae, Emesinae, true bugs, Ploiaria, Bargylia, taxonomy, Australia

Introduction The thread-legged assassin bugs (Hemiptera: Heteroptera: Reduviidae: Emesinae) of Australia were revised by WYGODZINSKY (1956); the Australian species were addressed in a monograph on the world fauna (WYGODZINSKY 1966) as well. Only WYGODZINSKY (1974), RÉDEI (2007), TATARNIC & CASSIS (2011), TATARNIC et al. (2011), and TATARNIC (2013) have provided additional data on the fauna of the continent (including its territorial islands). The subject of the current contribution is the description of two new species, one in genus Ploiaria Scopoli, 1786 and the other in Bargylia Stål, 1866, from mainland Australia. Ploiaria is a cosmopolitan genus with about 130 species worldwide. WYGODZINSKY (1956) revised the Australian fauna and described the majority of its endemic species. RÉDEI (2007) described an additional species. Currently there are eleven species known from Australia, two of them cosmopolitan, the rest endemic. One new species is described in the present paper. Bargylia is an endemic Australian genus. It was redefined, its type species described and fixed as type species by WYGODZINSKY (1951). Three additional new species were described and members of the genus were keyed by WYGODZINSKY (1956). The genus was redefined and redescribed, one species was removed, another was described as new, and the included species were revised and keyed once more by WYGODZINSKY (1966). The relationships and diagnostic characters of the genus and species groups within the genus were discussed and an additional species described by RÉDEI (2007). The genus now contains five species, and a further one is described in the present paper.

347 D. RÉDEI

This contribution is dedicated to two eminent Czech hemipterists, Jaroslav L. Stehlík and Pavel Lauterer, both of whom worked for the Moravian Museum for many years. With this paper I would like not only to pay tribute to their scientific contributions but also to acknowledge the great improvements they have made to the Hemiptera collection of the Hungarian Natural History Museum by identifying hundreds of pyrrhocoroid and psyllid specimens.

Material and methods Examinations of external structures were carried out using a Zeiss Discovery V8 stereoscopic microscope. Drawings were made with the aid of a camera lucida. Male genitalia were dissected after careful heating in hypertonic KOH solution, stained with Chlorazole Black E where necessary, and examined under a transmitted light microscope (Olympus CX21). Measurements were taken with a micrometer eyepiece.

Abbreviations for depositories: HNHM ...... Hungarian Natural History Museum, Budapest, Hungary NHRS ...... Naturhistoriska Riksmuseet, , Sweden

Taxonomy Genus Ploiaria Scopoli, 1786 Ploiaria Scopoli, 1786: 60. Type species: Ploiaria domestica Scopoli, 1786, by monotypy. For a complete list of synonyms of the genus see WYGODZINSKY (1966: 158), MALDONADO CAPRILES (1990: 108), PUTSHKOV & PUTSHKOV (1996: 158) and CASSIS & GROSS (1995: 292). A selected bibliography of the most important references on the genus was provided by RÉDEI & TSAI (2010: 21).

Selected references for the Australian fauna. WYGODZINSKY (1956): 215 (revision and key to Australian species), WYGODZINSKY (1966): 158 (redescription, revision and keys to world species), CASSIS & GROSS (1995): 292 (catalogue, Australia), MALDONADO CAPRILES (1990): 108 (catalogue, world), RÉDEI (2007): 365 (description of a new species), TATARNIC & CASSIS (2011): 22, 28 (fauna of Lord Howe and Norfolk Islands, description of a new species). Distribution and diversity. Ploiaria is a cosmopolitan genus containing about 130 species. Eleven species are known from continental Australia, and an additional 13 species occur in the surrounding islands usually considered to belong to the Australian Region: New Guinea, New Zealand and the oceanic islands of the tropical Pacific Ocean (WYGODZINSKY 1956, 1966; RÉDEI 2007; TATARNIC & CASSIS 2011).

Ploiaria stehliki sp.nov. (Figs 1–16)

Type material. Holotype: ♂, Australia: [1] “Derby” [printed], [2] “N. V. Austr. \ Mjöberg” [printed], [3] “okt.” [printed], [4] “REDUVIIDAE [handwritten] \ Emesinae [handwritten] \ Genn.? sp.? [handwritten] \ Froeschner [printed] 69 [handwritten]”; pinned, segments III–IV of left and segments II–IV of right antenna lacking (deposited in NHRS, identifier: NHRS-GULI000006461). Paratypes: with labels identical with those of [1], [2] and [3] of holotype (5 ♂♂ 4 ♀♀ deposited in NHRS, identifiers: NHRS-GULI000006462–6467, 6481, 6917–6918; 1 ♂ deposited in HNHM).

348 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of thread-legged assassin bugs (Reduviidae: Emesinae)

Diagnosis. Medium-sized species of Ploiaria (body length 10–11 mm) with quite uniformly brown ground colour, lacking contrasting pattern elements; known only from macropterous specimens of both sexes. It may be recognized by the following combination of characters: postocular portion of head much shorter than length of eye (Figs 1–2), posteriorly truncate in dorsal view (Fig. 1); pronotum about 1.1–1.2 times as long as its greatest width (Fig. 1); fore femur relatively slender (about seven times longer than its greatest diameter), ventral surface with a small, angulate tumescence at its extreme base (Fig. 5: arrow), anteroventral and posteroventral series of simple, relatively narrow spines inserted on minute, wart-like basal tubercles (Figs 4–5); genital capsule (Figs 7–8) relatively short, laterally flattened, with a relatively short and narrow, spine- like median process projecting posterodorsad. Description. Macropterous male and female. Colour, integument and vestiture. General colour brown, with paler areas but without distinct contrasting pattern; head brown, a pair of submedian longitudinal fasciae on anterior lobe, converging posteriorly, dark brown, ventral surface of head yellowish- brown; antenna dark brown; labium yellowish-brown; prothorax brown, dorsally marked with a fine, longitudinal-median yellowish line; yellowish-brown ventrally; fore leg quite uniformly yellowish-brown; mesonotum brown, its median longitudinal impression yellowish, membranous suture between mesonotum and mesopleuron whitish; mesoscutellum brown; fore wing brown; dorsal portion of pterothoracic pleuron brown, ventral portion of pterothoracic pleuron, whole of pterothoracic sternum and dorsal and ventral surfaces of abdomen yellowish-brown; mid- and hind coxae and trochanters yellowish-brown, femora brown with a pale, narrow annulus apically, tibiae dark brown, gradually becoming lighter towards apex, with a pale, narrow annulus basally. Integument dull, delicately granulate, body and legs covered in very short, fine, inconspicuous hairs, appears glabrous even when observed under a stereoscopic microscope at 50× magnification; antenna without conspicuous hairs in both sexes. Structure. Head (Figs 1–2) 1.1–1.2 times as long as its width across eyes, 2.5–2.6 times as broad as interocular distance; preocular portion about 2.2 times longer than postocular portion, relatively narrow and subparallel forward of eyes; interocular furrow weakly curved posteriad; postocular portion about 1.3 times as wide as preocular portion, broadly rounded, posteriorly truncate in dorsal view; eyes large, semiglobular, strongly protruding laterally in both sexes, approaching but not surpassing ventral outline of head in lateral view; first visible segment of labium slightly longer than second visible segment (and about 1.25 times as long); antennal insertions situated slightly below centre of preocular portion. Pronotum (Figs 1–2) short, 1.1–1.2 times as long as its greatest width (somewhat forward of its mid-point), anterior lobe broadly rounded laterally, with a fine furrow along midline, posterior lobe short, collar-like, distinctly broadened posteriorly. Pterothorax. Mesonotum 1.3–1.4 times longer than pronotum, 1.2–1.3 times longer than its basal width; metasternum with a fine, narrow, median longitudinal carina. Fore wing (Fig. 3) elongate; r-m cross vein very short, situated slightly basad to the cross vein between Cu and AA1+2; cubital cell (= discal cell sensu WYGODZINSKY 1966; Fig. 3: cuc)

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Figs 1–6. Ploiaria stehliki sp.nov. 1 – head and anterior portion of thorax, dorsal view; 2 – same, lateral view; 3 – right fore wing; 4 – right fore leg, anterior (= inner) view; 5 – trochanter and base of femur of right fore leg, anterior (= inner) view; 6 – pretarsus of right fore leg, posterior (= outer) view. Lettering: ac = anterior claw; pc = posterior claw; for fore wing venation, see the text. Scales in mm.

nearly two times as long as apical free portion of M, about as long and as broad as length and greatest diameter of the area delimited by Sc+R, M, r-m and Rs cross-veins respectively; apical free portion of M running at about same distance from anterior and posterior margins of wing; Rs joined to apical free portion of M around midway between

350 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of thread-legged assassin bugs (Reduviidae: Emesinae) apex of cubital cell and apex of wing. Fore leg (Fig. 4) relatively slender; coxa 1.6–1.7 times as long as pronotum; trochanter with two narrow spines inserted on minute, wart- like basal tubercles and with a few additional setae; femur relatively slender, 6.9–7.05 times longer than its greatest diameter (in its basal third), dorsal and ventral outlines nearly subparallel, anteroventral and posteroventral series of simple, relatively thin spines inserted on minute, wart-like basal tubercles; anteroventral series of about 40 spines, posteroventral series of about 55–60 spines, anteroventral series uninterrupted at base; ventral surface of fore femur with a small, angulate tumescence at its extreme base (Fig. 5: arrow), basal fifth of ventral surface with 10–15 spines somewhat longer and stronger than remaining spines of posteroventral series, arranged in two or three irregular rows; tibia slender, straight, about 0.6 times as long as femur, ventral surface with a single series of deflected spine-like setae ventrally; tarsus slender, straight, about 0.55 times as long as tibia, with two series of short, fine, adpressed spine-like setae ventrally, with two claws of different sizes, posterior (= outer) claw (Fig. 6: pc) shorter and stouter than anterior (= inner) claw (Fig. 6: ac). Mid- and hind legs delicate, hind femur about as long as body from apex of head to apex of abdomen.

Abdomen relatively slender; posterior margin of sternite VII of female (Fig. 16: s 7) slightly rounded posteriorly. External male genitalia. Genital capsule (Figs 7–8) relatively short, laterally flattened, anterior and posterior apertures separated by a broad dorsal sclerotized, bridge- like portion, so the posterior aperture occupies only about the posterior half of the segment, its proximal margin broadly rounded and broadly emarginate in middle third; cup-like sclerite with a relatively short, narrow, spine-like median process projecting posterodorsad, gradually narrowing towards apex; genital capsule tumescent at its posterior margin below median process. Parameres (Figs 9–11) sickle-shaped, with long, scattered setae. Phallus (Figs 12–13): articulatory apparatus (Fig. 13: aa) restricted to extreme base of phallus, basal plates (Figs 12–13: bp) subparallel in their proximal halves, diverging in their distal halves; phallosoma (Fig. 13: phs) simple, tubular, curved; basal aula with a pair of distinct lobes (Figs 12–13: lba) proximally; “struts” (Figs 12–13: str) separate proximally, sharply converging towards midline and fused quite close to centre, fused portion gradually narrowing towards distal portion and fused with dorsal sclerotized plate of phallosoma, pointed distally; dorsal sclerotized plate (Figs 12–13: dsp) narrowly split along midline; ventral sclerotized plate (pedicel?) (Figs 12–13: vsp) running along whole length of phallotheca; endosoma with densely-packed, denticle-like sclerites (Fig. 13: ses).

External female genitalia (Figs 14–16). Tergite VIII (Figs 14–15: t8) broadly triangularly produced posteriorly at middle; valvifers VIII (Figs 14–16: vf8) broadly surrounding derivatives of segments IX laterally, their inner margins straight, adjacent along midline, with a long, narrow apophysis projecting into segment VII (Fig. 15: avf8); valvula VIII (Figs 14, 16: va8) short, simple, obliquely directed; tergite IX (Figs 14–15: t9) broadly emarginate basally, subtruncate apically in dorsal view; valvifer IX (Fig. 14: vf9) and valvula IX (Fig. 14: va9) both rod-shaped, simple; gonoplac (Figs 14–16: gpc) short, directed posteriad, contralateral gonoplacs not fused.

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Figs 7–14. Ploiaria stehliki sp.nov. 7 – genital capsule, dorsal view; 8 – same, lateral view; 9–10 – left paramere, two different aspects; 11 – apical portion of left paramere; 12 – phallus, dorsal view; 13 – same, lateral view; 14 – apex of abdomen of female, lateral view. Arrow in Fig. 10 shows aspect of Fig. 11. Lettering: aa = articulatory apparatus; dsp = dorsal sclerotized plate; gpc = gonoplac; lba = lobe of basal aula; phs = phallosoma; s7 = sternite VII; ses = denticle-like sclerites of endosoma; str = “struts”; t8–t9 = tergites VIII and IX; va8–va9 = valvulae VIII and IX; vf8–vf9 = valvifers VIII and IX; vsp = ventral sclerotized plate; X = segment X; XI = segment XI. Scales in mm.

Measurements (in mm) (holotype ♂ / paratypes, n = 3 ♂♂ 3 ♀♀). Body length 10.7 / 10.0–11.0, length of head 1.19 / 1.24–1.29, width across eyes 1.10 / 1.05–1.10, interocular distance 0.42 / 0.41–0.42, lengths of antennal segments I : II as 6.88 / 6.40–6.80 : 5.94 / 5.50–5.90 (segments III–IV lacking in all specimens), median length of pronotum 1.06 / 1.07–1.17, greatest width (across anterior lobe) 0.97 / 0.91–0.98,

352 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of thread-legged assassin bugs (Reduviidae: Emesinae)

Figs 15–16. Ploiaria stehliki sp.nov. 15 – apex of abdomen of female, dorsal view; 16 – same, ventral view.

Lettering: avf8 = apophysis of valvifer VIII; t7 = tergite VII; others as in Fig. 14. Scale in mm.

length of fore coxa 1.76 / 1.78–1.98, length of fore femur 3.28 / 3.15–3.53, greatest diameter 0.47 / 0.45–0.51, length of fore tibia 2.05 / 1.86–2.08, length of fore tarsus 1.10 / 1.05–1.22, median length of mesonotum 1.60 / 1.34–1.55, posterior width 1.22 / 1.19–1.20, length of fore wing 7.15 / 7.10–7.50, greatest width 1.60 / 1.55–1.60, greatest width of abdomen 7.15 / 7.10–7.50. Etymology. Patronymic, dedicated to Jaroslav L. Stehlík for the occasion of his 90th birthday, in recognition of his achievements in the taxonomy, faunistics and bionomics of various groups of Heteroptera. Distribution. Australia: Western Australia. Discussion. Ploiaria stehliki sp.nov. is morphologically not notably similar to any of the described congeners from continental Australia, so its taxonomic relationships are uncertain. The distinct tumescence at the extreme base of the ventral surface of the fore femur (Fig. 5: arrow) is a peculiar autapomorphy of this species. I am not aware of any other species of Ploiaria exhibiting this character. The general habitus of this new species seems more or less similar to that of P. weiri Tatarnic & Cassis, 2011, described from Norfolk Island, although the rather superficial description of the latter species, ignoring several taxonomically important characters, renders comparison difficult. Some important differences between the two species (shape of postocular portion of head, base of ventral surface of fore femur, fore wing venation) makes it probable that, in spite of their superficial similarity, the two species are not phylogenetically close.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 353 D. RÉDEI

Figs 17–20. Bargylia lautereri sp.nov. 17 – head, thorax and first abdominal segment, dorsal view; 18 – same, lateral view; 19 – abdomen of female, dorsal view; 20 – same, lateral view. Scales in mm.

354 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of thread-legged assassin bugs (Reduviidae: Emesinae)

Figs 21–26. Bargylia lautereri sp.nov. 21 – right fore leg, anterior (= inner) view; 22 – apex of abdomen of male, lateral view; 23 – same, dorsal view; 24 – genital capsule, dorsal view; 25–26 – left paramere, two

different aspects. Lettering: s7 = sternite VII; t7 = tergite VII; VIII = segment VIII. Scales in mm.

Genus Bargylia Stål, 1866

Bargylia Stål, 1866a: 163. Type species: iuncea (non Erichson, 1842): STÅL (1874: 96), misidentification (= Bargylia stali Wygodzinsky, 1951), by subsequent designation by WYGODZINSKY (1951): 611, 616 (see Nomenclature paragraph below).

References. STÅL (1866b): 168 (listed), STÅL (1874): 94 (in key), 96 (catalogue), LETHIERRY & SEVERIN (1896): 75 (catalogue), WYGODZINSKY (1951): 610 (designation of type species, redescription), WYGODZINSKY (1956):

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196 (in key), 197 (identity, taxonomic history, nomenclature, key to species), WYGODZINSKY (1966): 434 (in key), 449 (redescription, revision, key to species), MALDONADO CAPRILES (1990): 121 (catalogue), KERZHNER (1992): 49 (type species, nomenclature), CASSIS & GROSS (1995): 296 (catalogue), RÉDEI (2007): 372 (diagnostic characters, species groups).

Nomenclature. WYGODZINSKY (1951) demonstrated that the genus Bargylia was based on a misidentified type species, and he selected the taxonomic species actually involved in the misidentification (B. stali Wygodzinsky, 1951) as type species of the genus. In this, and also in a subsequent paper (WYGODZINSKY 1966: 449), he indicated that an application had been submitted to the International Commission on Zoological Nomenclature intending to fix B. stali as type species of Bargylia. KERZHNER (1992), referring to WYGODZINSKY (1966), also mentioned that such an application had been submitted. However, it appears that no such Case or Opinion has been published to date. The current edition of the International Code of Zoological Nomenclature (4th edition, ICZN 1999, Art. 70.3) does not require such cases to be referred to the commission but requires a First Reviser Act to have the type species fixed. Although the condition “the author must refer to this [= 70.3] Article” is of course not met, in all other respect WYGODZINSKY’s (1951) First Reviser Act fully conforms to Art. 70.3.2 and was uniformly followed by all subsequent authors; it is therefore here considered as a valid type species fixation. Distribution and diversity. Five species of Bargylia were known prior to this study, all occurring in Australia (RÉDEI 2007). An additional new species is described below.

Bargylia lautereri sp.nov. (Figs 17–32)

Type material. Holotype: ♂, Australia: [1] “Cook- \ town” [printed], [2] “Queensl. \ Mjöberg” [printed], [3] “REDUVIIDAE [handwritten] \ Emesinae [handwritten] \ Genn.? spp.? [handwritten] \ Froeschner [printed] 67 [handwritten]”; mounted on pointed card, left antennal segment IV and both mid tibiae and tarsi lacking (deposited in NHRS, identifier: NHRS-GULI000006482). Paratypes: with labels identical with those of [1] and [2] of holotype (1 ♂ 5 ♀♀ and 1 specimen lacking abdomen deposited in NHRS, identifiers: NHRS- GULI000006483–6488 and 6919; 1 ♀ deposited in HNHM). Diagnosis. A small member of the genus Bargylia (body length 8–9 mm) diagnosed by the following combination of characters: body surface finely but distinctly granulate; thoracic segments relatively short; clypeus and antenniferous tubercles without spine-like process (Figs 17–18); fore femur relatively stout (9–9.7 times as long as its greatest diameter), with long spiniferous processes, 5 processes of posteroventral series subequal to or longer than diameter of segment (Fig. 27). Description. Apterous male and female. Colour, integument and vestiture. Body with complex pattern of dark brown to yellowish areas (Figs 17–18); head yellowish, a pair of lateral longitudinal fasciae as broad as height of eye running from base of head to antenniferous tubercles, a pair of more narrow submedian longitudinal fasciae running from posterior fourth of postocular portion to antenniferous tubercles, and gena dark brown; antenna brown; labium yellowish-brown; prothorax yellowish, broadly dark brown laterally; fore leg yellowish,

356 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of thread-legged assassin bugs (Reduviidae: Emesinae)

Figs 27–32. Bargylia lautereri sp.nov. 27 – phallus, ventral view; 28 – same, lateral view; 29 – basal plate; 30 – apex of abdomen of female, dorsal view; 31 – same, lateral view; 32 – same, ventral view. Lettering: ism9–10 = intersegmental membrane between segments IX and X; phm = phallosoma mouth; other abbreviations as in Figs 12–16. Scales in mm.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 357 D. RÉDEI posterior surface of femur and apical third of tibia more or less distinctly suffused with brown; median thirds of meso- and metanota yellowish, lateral thirds brown; meso- and metathoracic pleurosternites (= fused pleurites and sternites) laterally brown, ventrally yellow; mid and hind coxae and trochanters yellowish, femora light brown, tibiae somewhat darker brown, with a narrow and rather indistinct pale annulus on femora subapically and tibiae sub-basally (usually more distinct on tibiae, frequently virtually lacking on femora); abdominal tergites yellowish to light brown, mottled with brown, most abdominal sternites yellowish to light brown, broadly suffused with brown along their lateral margins. Integument dull, delicately granulate, body appears glabrous. Structure. Head (Figs 17–18) about 1.6 times as long as its width across eyes, about 1.6 times as broad as interocular distance; portion behind antennifers nearly ovoid, with evenly convex gula, portion in front of antennifers strongly narrowing towards apex; clypeus without process; preocular portion about 1.6 times longer than postocular portion; postocular portion 1.15–1.2 times as wide as preocular portion, nearly globose; eyes small in both sexes, semiglobular, situated slightly closer to dorsal than to ventral surface of head in lateral view; first visible segment of labium surpassing level of antennifer but far distant from anterior margin of eye; antennal insertions situated slightly behind middle of preocular portion. Pronotum (Figs 17–18) moderately elongate, 3.0–3.1 times as long as its greatest width (in its anterior fourth), gradually narrowing posteriorly, posterior lobe short, collar- like, weakly demarcated. Pterothorax. Mesonotum 0.45–0.5 times as long as pronotum; metanotum 0.6–0.65 times as long as mesonotum. Fore leg (Fig. 21) relatively slender; coxa 0.95–1.0 times as long as pronotum; femur 1.5–1.6 times as long as coxa, relatively slender, 9.0–9.7 times as long as its greatest diameter (around middle); anteroventral series of about 30 spines inserted on elongate basal processes and an additional short spiniferous process next to basalmost process of posteroventral series; posteroventral series of 5–6 greatly elongate spiniferous processes (basal one longest, in some specimens nearly two times as long as diameter of femur) intermixed with about 15 shorter spiniferous processes; basalmost process about as long as, or slightly shorter than, distance from base of femur to insertion of that process; tibia about 0.35–0.4 times as long as femur; tarsus about 0.85 times as long as tibia. Abdomen (♀: Figs 19–20) slender, about 7.8 (♂) / 8.1–8.3 (♀) as long as its greatest diameter; spiracles I–VIII each situated on a large, conspicuous tubercle; posterior margin of tergite VII of male strongly produced towards the rear, covering genital capsule from above, segment VIII greatly exposed (Figs 22–23); posterior margin of tergite VII of female straight (Fig. 27: t7), that of sternite VII broadly and obtusely subtriangularly produced posteriad at middle (Fig. 32: s7). External male genitalia. Genital capsule (Fig. 24) laterally flattened, anterior and posterior apertures separated by a narrow, dorsal, sclerotized, bridge-like portion close to base of segment; apical portion of cuplike sclerite protruding nearly vertically from genital capsule, trapezoid, apically relatively narrowly truncate in posterior view. Parameres (Figs 25–26) simple, with straight basal portion and curved apical portion. Phallus (Figs 27–29) (homologies very unclear): basal plates (Figs 28–29: bp) robust,

358 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of thread-legged assassin bugs (Reduviidae: Emesinae) fused along their whole length except at extreme base, continued in a pair of elongate sclerites surrounding basal aula (basal portion of phallosoma) ventrolaterally (derivative of pedicel?), apical portion with a pair of rod-like processes surrounding phallotheca ventrally (derivatives of ventral sclerotized plate?); phallosoma (Fig. 28: phs) simple, membranous, phallosoma mouth (Fig. 28: phm) slightly widened. External female genitalia (Figs 14–16). Posterior margin of tergite VIII (Figs 30–31: t8) with a pair of long, broad, posteromedially-directed projections that converge towards midline and become contiguous distally; spiracles VIII large, situated on prominent protuberances; valvifers VIII (Figs 30–32: vf8) broadly surrounding derivatives of segments IX laterally, their inner margins adjacent proximally, strongly diverging distally; valvula VIII (Figs 31–32: va8) small, semicircular, with median margin nearly straight; tergite IX (Figs 31: t9) greatly covered by tergite VIII, valvifer IX (Figs 30–31: vf9) and valvula IX (Figs 30–31: va9) rod-shaped; gonoplac (Figs 31–32: gpc) short, directed posteriad, apex curved downwards, contralateral gonoplacs fused apically. Measurements (in mm) (holotype ♂ / paratype ♂, n = 1 / paratypes ♀♀, n = 3). Body length 8.2 / 8.5 / 8.8–9.0, length of head 0.89 / 0.91 / 0.94–0.98, width across eyes 0.55 / 0.56 / 0.57–0.58, interocular distance 0.34 / 0.35 / 0.35–0.38, lengths of antennal segments I : II : III : IV as 3.33 / 3.40 / 3.40–3.50 : 2.73 / 2.85 / 2.95–3.05 : 0.25 / ? / 0.22–0.25 : 0.76 / ? / 0.78–?, median length of pronotum 1.36 / 1.40 / 1.48–1.50, greatest width 0.43 / 0.47 / 0.49–0.50, length of fore coxa 1.34 / 1.33 / 1.39–1.44, length of fore femur 2.03 / 2.11 / 2.20–2.21, greatest diameter 0.22 / 0.22 / 0.24–0.26, length of fore tibia 0.76 / 0.78 / 0.84–0.85, length of fore tarsus 0.63 / 0.69 / 0.70–0.72, median length of mesonotum 0.63 / 0.74 / 0.70–0.74, median length of metanotum 0.41 / 0.42 / 0.45–0.49, length of abdomen 4.70 / 4.85 / 5.45–5.50, greatest width 0.60 / 0.60 / 0.66–0.68. Etymology. Patronymic, dedicated to Pavel Lauterer for the occasion of his 80th birthday, in recognition of his contributions to the taxonomy, faunistics and bionomics of psyllids and auchenorrhynchans. Distribution. Australia: Queensland. Discussion. RÉDEI (2007) highlighted problems of discrimination between the genera Bargylia and Bobba Bergroth, 1914 arising from the discovery of a species showing a combination of diagnostic characters from the two genera (B. longispina Rédei, 2007). In contrast with the last-named species, the new species B. lautereri sp.nov. is morphologically similar to the remaining four species of the genus and does not exhibit Bobba characters. Bargylia lautereri sp.nov. has a mosaic-like combination of the characters used by RÉDEI (2007) for defining a “stali-group” and a “longinota-group”, thus making separation of these two species groups impossible. It is similar to B. stali Wygodzinsky, 1951 and B. grossi Wygodzinsky, 1966 in the relatively stout fore femur (although its femur is somewhat more narrow than that of the two species) and the presence of long spiniferous processes on the fore femur, but it differs from these species in the absence of a spine-like projection on the clypeus, the conspicuously longer spiniferous processes

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 359 D. RÉDEI of the posteroventral series of the fore femur, and the relatively longer fore tarsus. It is similar to B. babinda Wygodzinsky, 1956 and B. longinota Wygodzinsky, 1956 in the absence of a spine-like projection on the clypeus, but these two species conspicuously differ from the new species in their smooth or inconspicuously granulate integument, more strongly elongate thoracic segments (particularly B. longinota), conspicuously more gracile fore leg, the fore femur being armed with shorter spiniferous processes, and the distance between the base of ventral surface of femur and lowest spiniferous process being distinctly larger than the length of the process. Finally, B. longispina Rédei, 2007 may easily be distinguished from the new species by its coarsely granulate body, produced antenniferous tubercle, and the widened abdomen of the male. The shape of tergite VIII (largely covering tergite IX, provided with a pair of long posterior projections) (Fig. 30: t8) seems unique in the genus, although no female of B. longispina has yet been published.

Acknowledgements I am grateful to Gunvi Lindberg (NHRS) for her hospitality during my visit to Stockholm and for access to the collection under her care; to Dimitri Forero (Pontifica Universidad Javeriana, Bogotá) for reviewing the manuscript; to Jing-Fu Tsai (Hokkaido University, Sapporo) for discussion about the homologies of the male genitalia; to András Orosz (HNHM) for various kinds of help; and to Petr Kment (National Museum, Prague) and Igor Malenovský (Moravian Museum, Brno) for their invitation to contribute to this volume and for their careful editorial work.

References

CASSIS G. & GROSS G. F. 1995: Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha). In: HOUSTON W. W. K. & MAYNARD G. V. (eds.): Zoological Catalogue of Australia. Vol. 27.3A. CSIRO Australia, Melbourne, xv + 506 pp. ICZN (INTERNATIONAL COMMISSON ON ZOOLOGICAL NOMENCLATURE) 1999: International Code of Zoological Nomenclature. Fourth edition. International Trust for Zoological Nomenclature, London, 306 pp. KERZHNER I. M. 1992: Nomenclatural and bibliographic corrections to J. Maldonado Capriles (1990) “Systematic catalogue of the Reduviidae of the world (Insecta: Heteroptera)”. Zoosystematica Rossica 1: 46–60. LETHIERRY L. & SEVERIN G. 1896: Catalogue général des Hémiptères. Tome III. Hétéroptères, , Phymatidae, , Hebridae, , Henicocephalidae, Reduviidae, , Apophilidae, , , . Friedländer & Fils, Berlin, 275 pp. MALDONADO CAPRILES J. 1990: Systematic catalogue of the Reduviidae of the world (Insecta: Heteroptera). Caribbean Journal of Sciences, Special Edition, x + 694 pp. PUTSHKOV P. V. & P UTSHKOV V. G. 1996: Family Reduviidae Latreille, 1807 – assassin-bugs. Pp. 390–538 In: AUKEMA B. & RIEGER CH. (eds): Catalogue of the Heteroptera of the Palaearctic Region, vol. 2. The Netherlands Entomological Society, Amsterdam, xiv + 361 pp. RÉDEI D. 2007: New and little-known thread-legged assassin bugs from Australia and New Guinea (Heteroptera: Reduviidae: Emesinae). Acta Zoologica Academiae Scientiarum Hungaricae 53(4): 363–379. RÉDEI D. & TSAI J. F. 2010: A survey of the emesine assassin bugs of the tribes Collartidini, Leistarchini, , and Metapterini of Taiwan (Hemiptera: Heteroptera: Reduviidae). Deutsche Entomologische Zeitschrift 57(1): 11–36.

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SCOPOLI J. A. 1786: Deliciae faunae et florae insubricae, seu novae aut minus cognitae species plantarum et animalium quas in Insubria austriaca tum spontaceas quum exoticas vidit descripsit et aeri incindi curavit. Vol. 1. Salvator, Ticini, ix + 85 pp. STÅL C. 1866a: Hemiptera africana. Vol. 3. Norstedtiana, Holmiae, [1865], 200 pp. STÅL C. 1866b: Analecta hemipterologica. Berliner Entomologische Zeitschrift 10: 151–172. STÅL C. 1874: Enumeratio Hemipterorum. Bidrag till en Förteckning öfver alla hittills kända Hemiptera, jemte systematiska meddelanden. 4. Kongliga Svenska Vetenskaps-Akademiens Handlingar 12(1): 1–186. TATARNIC N. J. 2013: Mafulemesa schuhi (Heteroptera: Reduviidae: Emesinae), a new species from Australia. Entomologica Americana 118(1–4) [2012]: 274–277. TATARNIC N. J. & CASSIS G. 2011: The thread-legged bugs (Hemiptera: Heteroptera: Reduviidae: Emesinae) of Lord Howe and Norfolk Islands. Zootaxa 2967: 21–43. TATARNIC N. J., WALL M. A. & CASSIS G. 2011: A systematic revision of the Australian ploiarioline thread- legged assassin bugs (Hemiptera: Reduviidae: Emesinae). Zootaxa 2762: 1–30. WYGODZINSKY P. W. 1951: On Bargylia Stål, 1865, Pseudobargylia, gen. nov. and Leistarches serripes Dohrn, 1860 (Emesinae, Reduviidae: Hemiptera). Annals and Magazine of Natural History, Series 12 4(42): 609–617. WYGODZINSKY P. W. 1956: Synopsis of the Australian Emesinae (Hemiptera: Reduviidae). University of California Publications in Entomology 11(4): 193–246. WYGODZINSKY P. W. 1966: A monograph of the Emesinae (Reduviidae, Hemiptera). Bulletin of the American Museum of Natural History 133: 1–614, plates 1–4. WYGODZINSKY P. W. 1974: Description of the first winged species of Pseudobargylia (Emesinae, Reduviidae, Hemiptera). Memoirs of the National Museum of Victoria 35: 111–113.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 363–379, 2013

Two new genera and four new species of micropterous Aradidae from Malaysia (Hemiptera: Heteroptera)

ERNST HEISS1 & PETR BAÒAØ2 1 Tiroler Landesmuseum, Josef-Schraffl-Strasse 2a, A-6020 Innsbruck, Austria; e-mail: [email protected] 2 Moravian Museum, Department of Entomology, Hviezdoslavova 29a, Brno, CZ-627 00, Czech Republic; e-mail: [email protected]

HEISS E. & BAÒAØ P. 2013: Two new genera and four new species of micropterous Aradidae from Malaysia (Hemiptera: Heteroptera). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 363–379. – The following new genera and species of Aradidae (Heteroptera) are described and figured: Carventinae, Langkawiaptera gen.nov. with L. spinosa sp. nov from Langkawi Island in Malaysia; and Mezirinae, Stehlikiessa gen.nov. with S. kmentiana sp.nov. from the Cameron Highlands in West Malaysia, and Smetanacoris parallelus sp.nov. and Smetanacoris lautereri sp.nov. from Sarawak in Borneo. A key to species of Smetanacoris is provided. Keywords. Hemiptera, Heteroptera, Aradidae, Carventinae, Mezirinae, taxonomy, new genera, new species, micropterous, Oriental Region, Malaysia

Introduction Sifting leaf litter or substrate under loose bark in subtropical and tropical forests in the Old World, a collection approach that is gaining in popularity, has led to the detection of a great number of new flat bug taxa (e.g. HEISS 1997, 2001, 2011; HEISS et al. 2012; HEISS & BAÒAØ 2013a, b, c). Most of them are particularly adapted to these nourishment- rich habitats and are represented by both apterous and micropterous forms. Their distribution range, however, is very limited and so endemism is to be expected. From material originating from the Malay peninsula and Sarawak province in Borneo, two new genera and four new species have been recognized and they are described and figured herein. We dedicate our paper to Dr. Jaroslav Stehlík on the occasion of his 90th birthday, in recognition of his friendship and outstanding contributions to our science and to our esteemed colleague Dr. Pavel Lauterer who is celebrating his 80th birthday.

Material and methods The specimens studied were partly borrowed from the Muséum d’histoire naturelle de la Ville de Genève or are from the collection of one of the authors (EH).

363 E. HEISS & P. BAÒAØ

Types are deposited in, or are held by: EHIA ...... Ernst Heiss collection, at Tiroler Landesmuseum, Innsbruck, Austria MHNG ...... Muséum d’histoire naturelle de la Ville de Genève, Geneva, Switzerland MMBC ...... Moravian Museum, Brno, Czech Republic NMPC ...... National Museum, Prague, Czech Republic

The usual incrustation that tends to obscure the body structures of litter-dwelling specimens was removed by soaking them in 10% KOH and rinsing in distilled water. Measurements were taken with a micrometer eyepiece and are given in millimetres. When citing the text on the labels attached on the same pin as the specimens, a forward slash (/) separates the lines of one label, and a double slash (//) separates different labels. The following abbreviations are used in the descriptions: deltg = dorsal external laterotergite (connexivum), mtg = mediotergite, ptg = paratergite, vltg = ventral laterotergite.

Taxonomy Subfamily Carventinae Usinger, 1950 Langkawiaptera gen.nov. Type species: Langkawiaptera spinosa sp.nov., here designated. Description. Apterous, body surface glabrous with pits and elevations and scattered incrustate patches of pilosity; colour dark brown to black, legs, antennae and lateral spines ochraceous yellow, pilosity stramineous, a dull, pale yellow. Head rectangular in outline with projecting clypeus reaching about half of antennal segment I; antenniferous lobes not developed; antennae shorter than width of head, segment I longest and thickest, bent at base, II shortest, III thinner and IV fusiform; eyes directed dorsolaterally with a large preocular tubercle; postocular lobes deeply excavated anteriorly then rounded and converging posteriorly to a constricted neck; rostrum arising from a slit-like atrium, rostral groove deep, closed posteriorly, shorter than head. Pronotum strongly transverse, a median depression separating the ovate, laterally- raised sclerites with an anterolaterally-directed spine at apex; a deep furrow separates the pronotum from mesonotum. Mesonotum consists of a transverse median ridge connecting the laterally-raised sclerites and bears a spine at the apex; posterior margin straight and strongly raised, sloping anteriorly to a deep, transverse furrow. Metanotum fused to mtg I+II forming a trapezoidal sclerite; anterior margin raised with two median and 2 (1+1) lateral, backward-directed projections, disk sloping posteriorly to deep impressions, their rear margins straight, carinate and smooth. Abdomen. Tergal disk with concave surface and deep apodemal impressions; lateral margin with a lateral spine at fused deltg I+II and a lateral tubercle on deltg III–VII; surface of I+II and III–VI carinate midway, deltg II+III not fused; tergite VII medially raised for the reception of the pygophore; spiracles III–VII placed on tubercles visible from above, spiracle VIII on ptg VIII.

364 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genera and species of micropterous Aradidae

Figs 1–6. 1 – Langkawiaptera spinosa gen. et sp.nov., holotype, male, dorsal (3.75 mm). 2–3 – Stehlikiessa kmentiana gen. et sp.nov., holotype, male. 2 – dorsal view (2.55 mm); 3 – paratype, male, ventral view (2.55 mm). 4 – Smetanacoris apanius Heiss, 1989, paratype, female, dorsal view (2.42 mm). 5 – Smetanacoris parallelus sp. nov, paratype, female, dorsal view (2.5 mm). 6 – Smetanacoris lautereri sp.nov., paratype, female, dorsal view (2.35 mm).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 365 E. HEISS & P. BAÒAØ

Venter. Median part of pro-, meso- and metasternum smooth, sternites separated by deep, transverse furrows with the usual pattern of apodemal impressions. Legs unarmed, femora slightly incrassate, tibiae straight, claws with pulvilli. Etymology. Named after the Malaysian Island of Langkawi and the species’ apterous condition. Gender feminine. Differential diagnosis. The new genus is easily recognized and distinguished from all other apterous Carventinae recorded to date from Malay Peninsula and adjacent islands e.g.: Apteraradus Drake, 1957, Kiritshenkiana Kormilev, 1976, Morphocoris Kormilev, 1980, Cameronaptera Heiss, 2010a and Tiomanaptera Heiss, 2010b by body structure with prominent lateral spines on pro- and mesonotum and deltg I+II, head wide with short antennae and patches of velvet-like yellow pilosity on its surface. Included species. To date, the genus is monotypic, including only Langkawiaptera spinosa sp.nov. from Malaysia.

Langkawiaptera spinosa sp.nov. (Figs 1, 7–9) Type material. Holotype: male, labelled: ‘West Malaysia / Kedah, Pulau Langkawi / Gunung Raya, 700-800m / (6°23′N, 99°48′E) / 31 VIII 2004, leg. A. Schultz’ // ‘Holotype / Langkawiaptera gen.nov. / spinosa sp.nov. / des. E. Heiss & P. Baòaø 2013’ (MHNG). Description. Only an apterous male known. Surface of body glabrous with deep depressions and elevations, partly covered by patches of velvet-like incrustate pilosity (Figs 1, 7), colour dark brown to black, legs and antennae yellowish (Fig. 1). Head (Fig. 8) wider than long (0.9/0.8 mm); clypeus produced anteriorly with a dorsally-directed tubercle at apex, flanked by thin genae that diverge apically but are shorter than the clypeus and extend ventrally, leaving a median cleft for the ventrally- produced clypeus; antenniferous lobes not developed; antennae shorter than width of head (0.8/0.78 mm), segment I longest and thickest, bent at base, II shortest, III thinnest, IV fusiform with pilose apex; lengths of antennal segments I/II/III/IV = 0.25/0.15/0.19/0.20 mm; eyes granulate slightly stalked and directed dorso-laterally; large preocular tubercle present; postocular lobes excavated before eyes then rounded and produced laterally to outer level of eyes, converging posteriorly to a constricted neck; vertex carinate midway, flanked by 2 (1+1) ovate callosities at a lower level. Pronotum more than twice as wide as long (1.2/0.45 mm) with tufts of pilosity; collar ring-like, followed by a deep transverse depression at the rear that joins the transverse groove along the posterior margin; lateral sclerites raised toward lateral margin bearing a distinct spine on apex, anterolateral angles slightly produced over collar. Mesonotum five times as wide as long (1.5/0.3 mm); lateral sclerites raised with a distinct apical spine, medially connected by a ridge, lateral margins rounded; posterior margin straight and elevated, disk sloping anteriorly, then deeply depressed. Metanotum fused to mtg I+II, anterior margin highest with 2 median and 2 (1+1) lateral projections, surface sloping posteriorly with deep impressions before smooth, raised posterior margin.

366 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genera and species of micropterous Aradidae

Figs 7–9. Langkawiaptera spinosa gen. et sp.nov., holotype, male. 7 – dorsal view; 8 – head, dorsal view; 9 – tergal plate.

Abdomen (Fig. 9). Surface of tergal disk concave with distinct apodemal impressions; deltg I+II fused and triangular with a lateral spine antero- and posterolaterally; deltg I+II not fused to deltg III; lateral margin of deltg III–VII with a laterally-produced tubercle, surface with a longitudinal pilose carina; spiracles III–VII placed on a tubercle and visible from above, spiracles VIII on ptg VIII, pygophore conical, very wide, almost spherical from dorsal view, ptg VIII shorter than pygophore, apex rounded. Venter. Prosternum elevated midway with a deep pit; meso- and metasternum and following sternites II–VI smooth and lustrous at centre, lateral parts covered in velvet- like pilosity; sternites III–VII separated by transverse sutures; sternite VII with a median triangular elevation, matt, laterally with 2 (1+1) oval and velvet depressions. Legs unarmed, spine-like preapical bristles present on forelegs; femora moderately incrassate, tibiae cylindrical, tarsi 2-segmented with thin pulvilli. Measurements. Body length 3.75 mm (including the produced pygophore); antenna length 0.78 mm; ratio antenna length / head width = 0.87; abdomen width 1.85 mm. Differential diagnosis. The only species of this genus, it is easily recognized and distinguished from other apterous Carventinae by the characters provided in the generic description.

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Etymology. Named after the conspicuous lateral spines of the thorax; spinosus, -a, -um is a Latin adjective meaning spinose. Distribution. Endemic to Langkawi Island near the west coast of Malaysian Peninsula and not recorded elsewhere to date.

Subfamily Mezirinae Oshanin, 1908 Stehlikiessa gen.nov. Type species: Stehlikiessa kmentiana sp.nov., here designated. Description. Micropterous, small (2.5–2.9 mm); body stout with subparallel lateral margins, attenuated anteriorly; surface flat and submatt, pronotum with 4 (2+2) carinae. Head about as long as wide; clypeus short, reaching one-third length of antennal segment I; antenniferous lobes shorter than clypeus, apex acute; antennae about twice as long as width of head, segment I thickest, II shortest, III longest, IV with pilose apex, eyes inserted in head; postocular lobes granulate, converging posteriorly; rostrum arising from a slit-like atrium, as long as head, rostral groove open at the rear. Pronotum trapezoidal, lateral margins carinate converging forwards in angular fashion; sublateral raised carinae converging forwards in rounded fashion, joining raised anterior margin; disk sloping posteriad; larger tubercles are visible along lateral margin at a lower level. Mesonotum. Scutellum distinct, its surface flat; lateral wing pads with a longitudinal carina; canal of metathoracic scent gland visible from above, anterolateral to wing pads. Metanotum fused to mtg I+II, the fusion suture interrupted midway; surface smooth, posterior margin straight. Abdomen. Tergal plate flat with a carinate elevation medially on mtg V, apodemal impressions distinct; triangular fused deltg I+II with a longitudinal carina, not fused to deltg III, deltg III–VII separated by sutures, surface flat, lateral margins of deltg partly carinate, III, VI and VII in male and III–VII in female. Venter. Pro-, meso- and metasternum fused to sternites II+III, flattened at centre; sternites III–VII separated by deep furrows, surface smooth; metathoracic scent gland evaporatorium large, directed upwards, curved and raised with a slit-like ostiole. Legs unarmed, femora incrassate, tibiae cylindrical, preapical spine present, claws with thin pulvilli. Etymology. This interesting new genus is dedicated to our friend and mentor Jaroslav L. Stehlik from the Moravian Museum in Brno, an eminent scholar of the Pyrrhocoroidea. The gender is feminine. Differential diagnosis. The micropterous Mezirinae recorded to date from Malaysia belong to the genera Mastigocoris Matsuda et Usinger, 1957 (M. malayenis Kormilev, 1967 and M. truncatus Kormilev et Heiss, 1977), Kiritshenkiessa Kormilev, 1971 (K. spinipes Kormilev, 1971); to Hutanicoris Heiss, 1993 (H. carinatus Heiss, 1993); and to Pahangiessa Heiss, 1993 (P. bulboscutellata Heiss, 1993). Closest related to the species

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Figs 10–15. Stehlikiessa kmentiana gen. et sp.nov., paratype, male. 10 – dorsal view; 11 – ventral view; 12 – head, dorsal view; 13 – head, ventral view; 14 – thorax, dorsal view; 15 – thorax, ventral view.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 369 E. HEISS & P. BAÒAØ in hand appears to be Hutanicoris, sharing with the new genus its habitus and structure of pronotum but differing in the raised, not flat, scutellum with carinate lateral margins and in the ridge-like elevations marking the metanotum, lacking in Stehlikiessa gen.nov. Pahangiessa is larger (5.3–6.1 mm) and has a long clypeus, subrectangular head and a bulbously elevated scutellum; the micropterous form of Mastigocoris (macropterous morphs may occur in one population) lacks the carinate pronotum and has a triangular carinate scutellum, although it is of similar size and habitus, Kiritshenkiessa spinipes differs in its carinate scutellum and larger median elevation on the tergal plate and is also not considered for biogeographical reasons, since it is endemic to forests in south-west India. Included species. The genus is, at the moment, monotypic, including only Stehlikiessa kmentiana sp.nov. from Malaysia.

Stehlikiessa kmentiana sp.nov. (Figs 10–15) Type material. Holotype: male, labelled: ‘Malaysia. Pahang / Cameron Highlands / Gn. Beremban 1600-1800 m / 29.7.1993 leg. Schuh’ // ‘Holotype / Stehlikiessa gen.nov. / kmentiana sp.nov. / des. E.Heiss & P. Baòaø 2013’ (EHIA). Paratypes: 6 males, 1 female collected with holotype (EHIA, MMBC, NMPC); 2 males: ‘W Malaysia: Pahang / Cameron Highlands / trail 9, 1400m, 27.3.93 / Löbl & Calame, #21’ (MHNG); 1 male, 1 female: ‘AS-04/14 7 West Malaysia: Pahang, Frazer’s Hill / 2km south of town, 1200m / (3°41′N, 101°45′ E) / 28.VIII.2004, leg. A. Schultz / AS-04/12’ (MHNG); 1 male: ‘Malaysia. Pahang / Fraser’s Hill / 25 XI 1983 Heiss’ (EHIA). Description. Micropterous male (Figs 10–11), colour uniformly reddish-brown (Figs 2–3). Head (Figs 12–13) slightly wider than long (0.50/0.45 mm); clypeus elevated with a smooth, round subapical tubercle; antenniferous lobes shorter than clypeus, diverging anterolaterally with subacute apex; antennae 1.95 times as long as width of head, surface granulate, segment I thickest, constricted at base, II thinner and shortest, III longest, IV conical with pilose, truncate apex; length of antennal segments I/II/III/IV = 0.27/0.13/0.30/0.27 mm; eyes granulate, inserted in head; postocular lobes short and granulate, converging rearwards to constricted neck; vertex granulate at centre, flanked laterally by 2 (1+1) oval callosities; rostrum as long as head, arising from a slit-like atrium, rostral groove deep with carinate lateral margins, open posteriorly. Pronotum (Fig. 14) strongly transverse (1.1/0.35 mm); lateral carinate margins subparallel on posterior half then converging forwards in angular fashion, acute anterolateral angles produced over ring-like, medially-raised collar; 2(1+1) sublateral ridges curved anteriorly, there reaching the median elevation of collar; disk between these sublateral ridges concave and sloping to sinuate posterior margin with a central longitudinal groove; a few larger tubercles on a lower level of the lateral face of pronotum are visible from above. Mesonotum (Fig. 14). Scutellum flat at centre, anterolaterally acute angles slightly raised, posterior margin straight; lateral wing pads distinct, surface with a longitudinal carina; canal of metathoracic scent gland visible from above at anterolateral angle of wing pads.

370 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genera and species of micropterous Aradidae

Metanotum fused to mtg I+II (Fig. 14), surface smooth at centre, rugose laterally; fusion line marked by a thin suture interrupted midway; posterior margin straight. Abdomen. Tergal plate slightly concave with feeble apodemal impressions, a moderately median elevation is developed on mtg V; lateral margins seemingly doubled by reflexed vltg II–VII, which are visible from above as a small rim increasing in size from deltg II to VII; triangular deltg I+II fused, separated from deltg III by a suture, surface with a longitudinal ridge that continues on deltg III but is shorter and lower; surface of deltg III–VII flat, posterolateral margins of deltg VI and VII roundly elevated; tergite VII medially raised with a transverse granulate ridge; pygophore elongate, dorsal surface transversely rugose; paratergites VIII knob-like, reaching half of pygophore length. Venter. Prosternum anteriorly with 2 transverse carinae, then shallowly depressed and separated from mesosternum by a thin suture; meso- and metasternum flat at centre (Fig. 15), fused to sternites II+III (Fig. 11), the following sternites separated by transverse grooves; spiracles III–VI ventral on tubercles that form a “V” viewed from above, VII lateral and visible, VIII terminal on ptg VIII. Legs unarmed, beset with setigerous granulation; femora incrassate on apical half, trochanters distinct, tibiae straight with preapical spine on foreleg, claws with pulvilli. Female. Generally as male but of larger size; longitudinal ridges of deltg I+II continue along the outer margin of deltg III–VII, tergite VII truncate towards the rear, paratergites VIII reduced to 2 round, rearwards-produced tubercles bearing spiracles. Measurements. Holotype male: Body length 2.55 mm, antenna length 0.97 mm, abdomen width 1.25 mm. Paratype female: body length 2.9 mm, antenna length 1.1 mm, head width/length 0.55/0.52 mm, pronotum width/length 1.25/0.45 mm, abdomen width 1.5 mm; variation of body length in paratypes: males 2.50–2.60 mm, females 2.85–2.90 mm. Differential diagnosis. As this is to date the only species of this genus, it may clearly be recognized by the characters provided in the generic diagnosis and description. Etymology. It is our great pleasure to dedicate this new taxon to our friend Petr Kment (National Museum, Prague) in recognition of his important contributions to taxonomy and the faunistics of the Heteroptera, and of his successful engagement as editor of Acta Entomologica Musei Nationalis Pragae. The species name is designed as an adjective, kmentianus, -a, -um. Distribution. Recorded only from Cameron Highlands and adjacent Fraser’s Hill in Pahang Province, mainland Western Malaysia.

Smetanacoris Heiss, 1989 Smetanacoris Heiss, 1989: 2–5 (original description). Type species: S. apanius Heiss, 1989, by original designation. Notes. The micropterous Mezirinae genus Smetanacoris was erected for the species S. apanius Heiss, 1989 and S. sabahnus Heiss, 1989, both from the Malaysian province of Sabah in Borneo. Their most conspicuous character is the large, even dorsally expanded

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Figs 16–18. Smetanacoris parallelus sp.nov., holotype, male. 16 – total dorsal view; 17 – head, dorsal view; 18 – tergal plate.

evaporatorium of the metathoracic scent gland (HEISS 1989). Here we describe two additional new species from the Malaysian state of Sarawak, also in Borneo which also share this enlarged and extended evaporatorial surface. It is assumed to be an adaptation to the leaf-litter habitat, where it might be difficult to attract the opposite sex with gland secretions, or it may have a defence function that is facilitated by an increased evaporation surface. The two new species, S. parallelus sp.nov. and S. lautereri sp.nov. share a similar shape of the evaporatoria with laterally extended anterior part (Figs 24–26). The occurrence of clearly-distinguishable species from different localities of Sabah and Sarawak (Malayan part of Borneo) and the limited distribution range of flightless flat bugs indicate long, isolated evolutionary development resulting in endemic taxa.

Key to species of Smetanacoris Heiss, 1989 1(2) Evaporatorium of metathoracic scent gland developed posterolaterally to scent gland canal, its dorsal lobe short (Figs 21–23); scutellum with granulate elevation posteriorly (Figs 27–28)...... 3

372 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genera and species of micropterous Aradidae

Figs 19–20. Smetanacoris lautereri sp.nov., holotype, male. 19 – total dorsal view; 20 – abdomen, dorsal view.

2(1) Evaporatorium of metathoracic scent gland developed postero- and dorsolaterally to canal of scent gland, the dorsal lobe elongate and extending anterolaterally above the canal of the scent gland (Figs 24–26); scutellum with a narrow, apically-elevated ridge (Figs 29–30)...... 5 3(4) Median, backward-sloping depression between raised metanotal ridges smooth (Figs 4, 27)...... S. apanius Heiss, 1989 4(3) Median, backward-sloping depression between raised metanotal ridges with a median longitudinal carina and ovate depressions laterally (Fig. 28)...... S. sabahnus Heiss, 1989 5(6) Median, backward-sloping depression between raised metanotal ridges flat and smooth (Fig. 30); pronotum depressed at centre, delimited anteriorly by 2 (1+1) tubercles; outline of evaporatorium as in Figs 25–26; colour brown (Fig. 6)...... S. lautereri sp.nov. 6(5) Median, backward-sloping depression between raised metanotal ridges with a median longitudinal carina, the lateral depressions with transverse carinae and pits (Fig. 29); pronotum medially with a flat

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Figs 21–26. Smetanacoris species, evaporatorium of metathoracic scent gland, lateral view. 21–22 – S. apanius Heiss, 1989; 21 – holotype, male; 22 – paratype, female. 23 – S. sabahnus Heiss, 1989, holotype, male. 24 – S. parallelus sp.nov., holotype, male. 25–26 – S. lautereri sp.nov.; 25 – holotype, male; 26 – paratype, female. Scale bar 0.5 mm. Abbreviations (used in Figs 21 and 24): lm – lateral margin of mesonotum; mse – metathoracic scent gland evaporatorium; pr – lateral edge of pronotum; sgc – scent gland canal.

elevation, anterolaterally delimited by 2 (1+1) granulate ridges; outline of evaporatorium as in Fig. 24; colour blackish (Fig. 5)...... S. parallelus sp.nov.

Smetanacoris parallelus sp.nov. (Figs 5, 16–18, 24, 29) Material examined. Holotype: male, labelled: ‘E.Malaysia: Sarawak / confl. Suan Oyan and / Mujong riv. E Kapit / 150m, 19.V.1994 #6a / Löbl & Burckhardt’ // ‘Holotype / Smetanacoris / parallelus sp.nov. / des. E.Heiss & P. Baòaø 2013’(MHNG). Paratype: female, collected with holotype (EHIA). Description. Micropterous male, body subparallel (Figs 5, 16), surface with rugosities and carinae, dorsally visible evaporatoria elongate; colour blackish-brown. Head (Fig. 17) slightly wider than long (0.45/0.43); clypeus short with rounded apex and a shiny dorsal tubercle at centre; antenniferous lobes short and blunt with a dorsal

374 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New genera and species of micropterous Aradidae

Figs 27–30. Smetanacoris species, dorsal outline of scutellum and metanotum fused to mtgI+II, dorsal view. 27 – S. apanius Heiss, 1989; 28 – S. sabahnus Heiss, 1989; 29 – S. parallelus sp.nov.; 30 – S. lautereri sp.nov. Scale bar 0.5mm. Abbreviations: de – depression of mtg I+II; fa – flat sclerite of mtg I+II; gr – granulate elevation; mc – median carina; me – metanotal ridge; rs – ridge of scutellum; sc – scutellum; ts – transverse carinae.

tubercle; antennae 1.91 times as long as width of head, surface with setigerous granulation, segment I thickest, II shortest, III longest, IV with truncate, pilose apex; length of antennal segments I/II/III/IV = 0.22/0.12/0.27/0.24 mm; eyes inserted in head; postocular lobes angularly produced, then converging evenly to a narrow neck; vertex rugose; rostrum as long as head, arising from a slit-like atrium, rostral groove open posteriorly. Pronotum about twice as wide as long (0.75/0.4 mm); anterior margin raised midway with 2 (1+1) short, granulate ridges produced at the rear, followed at centre by lower subtriangular elevation; lateral sclerites elevated and granulate, separated from larger humeral tubercles and transverse carina of posterior margin by a sinuate groove.

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Mesonotum. Scutellum (Figs 16, 29) distinct, depressed at base, elevated posteriad, narrow median ridge highest at the rear, surface lateral to keel smooth, sloping to carinate lateral margins, anterolateral angles triangularly elevated; wing pads longer than scutellum with a median longitudinal ridge which is highest at the rear, laterally partly concealed by the elongate lobes of the evaporatorium of the metathoracic scent gland which is dorsally reflexed and visible from above; opening of metathoracic scent gland canal also visible in the middle of this evaporatorium (Fig. 16). Metanotum fused to mtg I+II forming 2 (1+1) inclined high lateral ridges, median depression between them with a carina at centre (Fig. 29), lateral parts with transverse carinae and pits, posterior margin nearly straight and carinate. Abdomen. Tergal plate (Fig. 18) rugose with deep impressions lateral of median ridge which is highest on mtg III; triangular fused deltg I+II separated from deltg III; surface of deltg III–VII raised with a median carina highest on deltg II and III with a posterolateral tubercle; tergite VII raised at centre with a knob-like apical tubercle, pygophore elongate with rugose surface, ptg VIII short and rounded. Venter. Pro-, meso- and metasternum and sternites II+III fused, prosternum elevated with an oval depression, meso- and metasternum flattened medially; evaporatorium of metathoracic scent gland large and trilobate (Fig. 24), extending posterolaterally and dorsally; spiracles III–VI ventral, VII lateral and visible from above, spiracle VIII terminal on ptg VIII. Legs unarmed, fore tibiae with a preapical spine-like comb, femora moderately incrassate midway, claws with pulvilli. Female. Generally as male, surface of deltg III–VII with 2 tubercles along inner margin and another one on posterolateral angle; tergite VII depressed at centre, raised to a transverse ridge posteriorly; ptg VIII consisting of two round tubercles connected by a thin, bridge-like sclerite, bearing spiracles at apex. Measurements. Holotype, male: body length 2.4 mm, antenna length 0.86 mm, abdomen width 0.95 mm. Paratype, female: body length 2.5 mm, antenna length 0.91 mm, ratio antenna length/head width = 2.02, head width/length 0.45/0.43 mm, pronotum width/length 0.85/0.40 mm, abdomen width 1.05 mm. Etymology. The species epithet is the Latin adjective parallelus, -a, -um, meaning parallel. The name refers to the subparallel body of this species. Differential diagnosis. Smetanacoris parallelus sp.nov. may be distinguished from S. apanius and S. sabahnus by its long evaporatorium with the dorsal lobe elongate and extending anterolaterally (short, not extending anterolaterally in S. apanius and S. sabahnus) and scutellum with a narrow, apically-elevated ridge (scutellum with a granulate elevation at the rear in S. apanius and S. sabahnus); from S. lautereri sp.nov. by median longitudinal carina on metanotum (metanotum flat in S. lautereri sp.nov.); by flat elevation on pronotum (pronotum with two tubercles in S. lautereri sp.nov.) and by shape of evaporatorium (see also Key). Distribution. Recorded to date only from central part of Sarawak, Kapit area.

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Smetanacoris lautereri sp.nov. (Figs 6, 19–20, 25–26, 30) Material examined. Holotype: male, labelled: ‘E. Malaysia: Sarawak / Gn. Matang, 20km W Kuching, 600m, 25 V 1994 / mix Dipterocarp forest / Löbl & Burckhardt # 11a’ // ‘Holotype / Smetanacoris / lautereri sp.nov. / des. E.Heiss & P. Baòaø 2013’ (MHNG). Paratype: female, ‘AS-EM 07/4 E.Malaysia / Sarawak, Gunung Gading / National Park near Lundu / primary forest, 600-800m / 01°42′50″N, 109°50′09″ E / 28 V 2007, leg.A.Schultz / AS-EM 07/4’ (EHIA). Description. Micropterous male (Figs 6, 19), colour uniformly brown (Fig. 6), surface submatt. As the basic structures are shared with S. parallelus sp.nov., the following description is abbreviated and refers primarily to differences between them to avoid or reduce unnecessary reiteration. Head slightly wider than long (0.47/0.43 mm); antenna 1.81 times as long as width of head, length of antennal segments I/II/III/IV = 0.20/0.11/0.27/0.27; other structures of head as in S. parallelus sp.nov. Pronotum twice as wide as long (0.75/0.35 mm); ring-like collar raised medially with 2 (1+1) round tubercles, depressed posteriad and sloping to convex transverse carina of posterior margin; lateral sclerites and humeral angles raised as in S. parallelus sp.nov. Mesonotum. Scutellum (Fig. 30) more than twice as wide as long, anterolateral angles triangularly elevated, disk raised to carinate posterior margin with a median ridge highest on posterior apex, lateral surface smooth; wing pads with a high longitudinal ridge which is lower midway; elongate lobes of metathoracic scent gland evaporatoria (Figs 25–26) partly covering lateral parts of wing pads. Metanotum with crescent-shaped ridges separated by a deep depression widening posteriad, its surface slightly concave and smooth. Abdomen. Surface of tergal plate smooth with a median elevation on mtg IV–VI and shallow lateral apodemal impressions; structure of deltg II–VII as in S. parallelus sp.nov., surface less rugose. Female. Generally as male in all essential structures. Measurements. Holotype, male: body length 2.4 mm, antenna length 0.86 mm, abdomen width 1.05 mm. Paratype, female: body length 2.35 mm, antenna length 0.8 mm, ratio antenna length / head width = 1.68, head width / length 0.23 / 0.24 mm, pronotum width / length 0.77 / 0.18 mm, abdomen width 1.1 mm. Etymology. Smetanacoris lautereri sp.nov. is dedicated to our colleague and friend Pavel Lauterer, a scholar of the Psylloidea and Auchenorrhyncha, on the occasion of his 80th birthday this year. Differential diagnosis. Smetanacoris lautereri sp.nov. can be distinguished from S. apanius and S. sabahnus by its long evaporatorium, dorsal lobe elongate and extending anterolaterally (short, not extending anterolaterally in S. apanius and S. sabahnus) and scutellum with a narrow, apically elevated ridge (scutellum with a granulate elevation at the rear in S. apanius and S. sabahnus). The new species differs from S. parallelus sp.nov. in its flat metanotum (median longitudinal carina on metanotum in S. parallelus sp.nov.); in two tubercles on pronotum (pronotum with flat elevation in S. parallelus sp.nov.) and in the shape of the evaporatorium (see also the Key). Distribution. This species is known only from east Sarawak.

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Acknowledgements The authors are grateful to Peter Schwendinger (Muséum d’histoire naturelle de la Ville de Genève) for the loan of interesting flat bug material and to Petr Kment (National Museum, Prague) and Lorène Marchal (Muséum national d’Histoire naturelle, Paris) for critical comments on the manuscript. We also thank Stefan Heim for taking the colour photographs. This paper appears through financial support provided to the Moravian Museum by the Ministry of Culture of the Czech Republic as part of its long-term conceptual development programme for research institutions (ref. MK000094862) and a grant of the Faculty of Science of Charles University in Prague (SVV-2013-267 201). Ernst Heiss would also like to express his particular thanks to his friend Rudolf Schuh (Wiener Neustadt) who collected new and rare Aradidae several times and donated them generously to his special aradid collection. Petr Baòaø would like also express his thanks to Peter Schwendinger, John A. Hollier and Ivan Löbl for their hospitality and help during his visit to Geneva.

References

DRAKE C. J. 1957: New apterous Aradids from the West Indies (Hemiptera). Philippine Journal of Science 85: 405–412. HEISS E. 1989: A new genus and two new species of micropterous Mezirinae from Sabah (Heteroptera, Aradidae). Entomofauna 10(1): 1–13. HEISS E. 1993: Two new genera of brachypterous Mezirinae from Malaysian rainforests (Heteroptera, Aradidae). Berichte des Naturwissenschaftlich-Medizinischen Vereins Innsbruck 80: 327–335. HEISS E. 1997: Two new genera of apterous Carventinae from New Guinea (Heteroptera, Aradidae). Mitteilungen der Münchner Entomologischen Gesellschaft 87: 73–79. HEISS E. 2001: Singhalaptera froeschneri gen. n., sp. n., from Sri Lanka (Heteroptera: Aradidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 53: 111–115. HEISS E. 2010a: Cameronaptera glabrinotum n.gen., n.sp. from Malaysia (Hemiptera: Heteroptera: Aradidae). Zootaxa 2530: 65–68. HEISS E. 2010b: A new genus and species of apterous Carventinae from Malaysia (Heteroptera, Aradidae). Denisia 29: 133–138. HEISS E. 2011: Ribesaptera elongata n. gen, n. sp., a curious apterous Mezirinae from Madagascar (Hemiptera: Heteroptera: Aradidae). Heteropterus Revista de Entomología 11(2): 273–277. HEISS E. & BAÒAØ P. 2013a: Ambohitantelya yuripopovi gen. nov. et sp. nov., a new apterous Mezirinae from Madagascar (Hemiptera: Heteroptera: Aradidae) with unique metathoracic evaporatoria. Zootaxa 3616(3): 291–297. HEISS E. & BAÒAØ P. 2013b: New apterous Carventinae from Sri Lanka and Southern India (Hemiptera: Heteroptera: Aradidae). Zootaxa 3647(3): 488–494. HEISS E. & BAÒAØ P. 2013c: Tiomanaptera schwendingeri, a new apterous carventine flat bug from Malaysia (Hemiptera: Heteroptera: Aradidae). Acta Entomologica Musei Nationalis Pragae 53: 33–38. HEISS E., BAÒAØ P. & RAHANITRINIAINA L. S. 2012: Two new species of the apterous Carventinae genus Comorocoris Heiss, 1985 from Madagascar (Hemiptera: Heteroptera: Aradidae). Zootaxa 3411: 63–68. KORMILEV N. A. 1967: Aradidae in the South Australian Museum, Adelaide II (Hemiptera, Heteroptera). Records of the South Australian Museum 15: 513–550. KORMILEV N. A. 1971: Mezirinae of the Oriental Region and South Pacific (Hemiptera-Heteroptera). Pacific Insects Monographs 26: 1–165. KORMILEV N. A. 1976: On the Aradidae from the Old and New World (Hemiptera, Heteroptera). Zoologica Scripta 5: 65–78. KORMILEV N. A. 1980: Homonymy in the Aradidae (Hemiptera). Pacific Insects 22: 328.

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KORMILEV N. A. & HEISS E. 1977: Five new Aradidae from the Oriental Region. Berichte des Naturwissenschaftlich-Medizinischen Vereins Innsbruck 64: 97–106. MATSUDA R. & USINGER R. L. 1957: Heteroptera: Aradidae. Insects of Micronesia 7: 117–172. OSHANIN B. F. 1908: Verzeichnis der palaearktischen Hemipteren mit besonderer Berücksichtigung ihrer Verteilung im Russischen Reiche. I. Band. Heteroptera. II. Lieferung. Tingidae – Acanthiidae. Ezhegodnik Zoologicheskago Muzeya Imperatorskoy Akademii Nauk 13(Suppl.): 395–586. USINGER R. L. 1950: The origin and distribution of apterous Aradidae. Pp. 174–179. In: Proceedings of the Eight International Congress of Entomology, Stockholm, August 9-14th, 1948. Axel B. Elfstroms Boktryckeri A.-B., 1030 pp.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 381–390, 2013

A new species of Dysdercus: Dysdercus stehliki sp.nov. (Hemiptera: Heteroptera: Pyrrhocoridae) from Brazil

CARL W. SCHAEFER Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269-3043, U.S.A.; e-mail: [email protected]

SCHAEFER C. W. 2013: A new species of Dysdercus: Dysdercus stehliki sp.nov. (Hemiptera: Heteroptera: Pyrrhocoridae) from Brazil. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 381–390. – A new species of the genus Dysdercus Guérin-Méneville, 1831, Dysdercus stehliki sp.nov., is named for Jaroslav L. Stehlík, the pre-eminent student of the superfamily Pyrrhocoroidea. Dysdercus stehliki sp.nov. is very close to D. longirostris Stål, 1861, with important similarities but also important differences. Dysdercus longirostris occurs near the coast of Brazil, and D. stehliki sp.nov. occurs inland, so far only from Viçosa, in the State of Minas Gerais, Brazil. The new species feeds on fallen fruit, especially Sterculia chicha A. St. Hil. (Malvaceae: Sterculioideae). Keywords. Hemiptera, Heteroptera, Pyrrhocoridae, Dysdercus, Sterculia, new species, host plant, Neotropical Region, Brazil

Introduction A year or so ago, colleagues at the Universidad Federal de Viçosa (Viçosa, State of Minas Gerais, Brazil) sent me some adults and nymphs of what I believed to be Dysdercus longirostris Stål, 1861. The bugs feed on fallen fruit of Sterculia chicha A. St. Hil. (Malvaceae: Sterculioideae), and we shall describe the immatures and their biology. But, looking more closely at the adults, I realized they were a new species and – in delight – I also learned that there would be a Festschrift for J. L. Stehlík. And so, ‘Nunc est bibendum, nunc pede libero pulsanda tellus…’ (Horace, Ode 37).

Material and methods The following abbreviations are used for type collections: CWS ...... Carl W. Schaefer personal collection, University of Connecticut, Storrs, USA; IOC ...... Instituto Oswaldo Cruz, Rio de Janeiro, Brazil; MMBC ...... Moravian Museum, Brno, Czech Republic; NMPC ...... National Museum, Prague, Czech Republic; USNM ...... National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A.

A slash (/) is used to divide rows of a single type label, a double slash (//) indicates different labels.

381 C. W. SCHAEFER

Taxonomy Dysdercus (Dysdercus) stehliki sp.nov. (Figs 1–4)

Type material. Holotype: ♂, ‘BRAZIL: Minas Gerais / State, Vicosa, June / 2011 / on Sterculia’ // ‘Coll. / Wagner de Souza Tavares’ (MMBC). Allotype ♀ (MMBC) and paratypes 7 ♂♂ and 1 ♀ (CWS, 1 ♂ 1 ♀; IOC, 2 ♂♂; MMBC, 1 ♂; NMPC, 1 ♂; USNM, 2 ♂♂), all with same labels. Description. Length (in mm): Holotype 9.75, allotype 14.43, paratypes (7 ♂♂) 9.62–13.13 (mean 11.53), paratype (1 ♀) 15.21. Head. Color: Deep brownish-red, but red around eyes, gula, and eyes themselves. Below, yellow or yellowish-red medially, red laterally; black or dark brown splotches laterally basally. In two specimens (2 ♂♂), head below yellow medially and sharply divided from red laterally; in another (allotype), yellow extending laterally, becoming fused with red. Antennae dark brown, often segment IV darker; proximal two-fifths of IV with white annulus. Rostrum light brown, segment IV darker or partly darker. Structure: Antennal segment I slightly curved, thicker than others, slight bulge distally, with a graduated dilation apically; II and III straight, narrower, no dilation; IV with a slight curvature, very slightly thicker; 4>2=1>3 (see Measurements, Table 1); all antennal segments with slight obscure pubescence, proximal part of III and all of IV with greater pubescence. First rostral segment slightly thickened, tapering apically; rostrum extending onto 6S (♂♂) or 7S (♀♀); 3≥2>1≥4 (see Measurements, Table 1). Thorax. Color: Corium and disc of pronotum light grayish-yellow, with gray fascia at base, and posteriorly with small strip of yellow at disc’s posterior edge; pronotal collar white; callus orange or orange-yellow; pre- and postcallar regions black or deep brown; raised lateral margin orange or orange-yellow. Scutellum orange, often with markings of gray, black, or dark brown midbasally and with a very thin line of black or dark brown laterally; sometimes apex somewhat more yellow. Membrane of wings deep brown, nearly black; with a very thin white or very pale brown line around corium. Prothoracic venter, anteriorly, white continuing with dorsal collar; all segments posteriorly and laterally white, with some pale yellow anteriorly; lateral to legs, orange, with a dark brown fascia anteriorly, and often with very small black spots laterally. Metathoracic scent gland paler yellow than surrounding areas, anterior region even sometimes paler. Legs (trochanter-tibia) brown, with dark brown spots; tarsus deeper brown, claws even deeper brown. Structure: Scutellum: slightly raised, sloping down laterally, basally, and apically; median depression mid-basally; sometimes more narrow than its width. Measurements of scutellum (in mm): holotype: length 1.58, width 1.58; allotype: length 1.75, width 1.72; paratypes (6 ♂♂ [one male obscured], 1 ♀): mean length 1.34, mean width 1.38 (range of lengths 0.99–1.55, of widths 0.97–1.65); length and width varying slightly, but in general of same length. Mesothoracic pleuron with some slight horizontal groovings (perhaps for metathoracic scent gland evaporatorium, whose base is quite close). Scent gland peritreme slightly less rough than surrounding metathorax. Abdomen. Color: Second sternum (2S) (1S not visible) to 5S, most of 3S, small area of 6S, 8S, and posterior half of 7S, yellow or slightly yellowish white; in darker specimens, dirty yellow (i.e., overtones of gray); rest of abdominal sterna orange-brown,

382 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Dysdercus stehliki sp.nov.

Figs 1–2. Dysdercus spp., habitus, dorsal view. 1 – D. stehliki sp.nov.: a – holotype, male; b – allotype, female. 2 – D. longirostris Stål, 1861, female.

or in darker specimens brown-black. In all specimens a thin, very dark brown stripe (from 3S to 7S) anteriorly. Exposed male and female genitalia orange-brown (if overall color darker, then much deeper brown). Structure: Laterally, very small dark brown pits, arranged in a series of concentric half-circles or half-triangles with slightly different texture; these pits of same color as abdominal sterna, behind and medial to spiracles and, on 5S–7S, anterior to the trichobothria (on 3S and 4S, trichobothria medial); pits of 2S–5S half-triangular (triangle pointing medially), 6S and 7S half-circular. Margins of 3rd to 5th sternites in females, and 3rd to 6th sternites in males, slightly explanate. Genital capsule (♂) with long hairs of same color as genital capsule itself, profusely on edge of ventral rim and edge of exposed cuplike sclerite (see terminology of SCHAEFER 1977, Table 1). Paramere laterally with large subapical spur and a smaller apical one twisted about 20° internally; between spurs, 0–4 very small teeth, these sometimes varying on parameres of same specimen (holotype lacks the teeth); paramere with sparse and short hairs on inner sides, especially its shank. Note: The hairs of the genital capsule occur in all Dysdercus males, but in Dysdercus stehliki sp.nov. they are longer and more profuse than I have seen in other males of other species. Differential diagnosis. The shorter length of this species, color differences, and the paucity of the paramere’s pilosity mark this species from its most closely related species, Dysdercus longirostris Stål, 1861. Additional differences are in the Discussion and Table 3.

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Figs 3–4. Dysdercus stehliki sp.nov., left paramere (small spots = bases of hairs): 3 – lateral view; 4 – medial view.

Etymology. I am wonderfully happy to name this species in honor of Jaroslav L. Stehlík, the pre-eminent student of Pyrrhocoroidea, who has done more to work out the speciation and the higher classification of this group than has any other person. From the 1960s, with studies on the Pyrrhocoroidea from E. S. Brown, and those from P. Basilewsky & N. Leleup, up until 2013 (and, we hope, later), he has worked on this group indefatigably, describing and analyzing both families (Largidae and Pyrrhocoridae) of the superfamily in all continents. Distribution. Brazil: Minas Gerais State.

384 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Dysdercus stehliki sp.nov.

Table 1. Adult antennal and rostral segment lengths of Dysdercus stehliki sp.nov.; range in parentheses (in mm). 1) Antennal segments of male paratypes: 1 specimen lacked antennae, 2 others were missing the fourth segment, and 1 other was missing the third.

Discussion The closest relative of Dysdercus stehliki sp.nov. is D. longirostris Stål, 1861. The latter occurs near the coast of central Brazil, from the states of Pernambuco through Espirito Santo to Rio de Janeiro and São Paulo. However, there may be some collecting bias, because many of the sites are near – or in – the city of Rio de Janeiro (see map, Fig. 74, of VA N DOESBURG (1968); and collecting sites, p. 126). The site of D. stehliki sp.nov. consists, now, only Viçosa, in Minas Gerais State; the city is inland, 300 km north of Rio de Janeiro and west of the coast by 150 km. I have looked at several other Brazilian sites, in my own collection and those borrowed; I have not seen D. stehliki sp.nov. elsewhere, but I have not made a major search. A significant feature of Dysdercus stehliki sp.nov. is the prolongation of the rostrum, which reaches to the sixth or slightly beyond (males) and well onto the seventh (females) abdominal sternum. In the great majority of New World Dysdercus, the rostrum reaches up to, or just beyond, the third abdominal sternum (3S). There appear to be only three other species of New World Dysdercus which also have a long rostrum: Dysdercus chiriquinus Distant, 1883, D. rusticus Stål, 1870, and D. longirostris. Dysdercus chiriquinus occurs in Central America and northern South America (northern Venezuela and northern Colombia). Its rostrum reaches 6S in the male and “end of abdomen in the female” (VAN DOESBURG 1968: 80). Dysdercus chiriquinus has a black spot on each corium, D. stehliki sp.nov. lacks the spot; D. chiriquinus also has a single spur on each paramere, D. stehliki sp.nov. has two. Dysdercus rusticus is distributed in east-central South America (including eastern Brazil) to Colombia (VAN DOESBURG 1968: 85). Like D. stehliki sp.nov., this species lacks

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 385 C. W. SCHAEFER any black on its corium; but its rostrum reaches onto 5S (male) or 6S (female), not to 6S and 7S. It also has a double spur on each paramere, but the spurs are of the same size; in D. stehliki sp.nov. the apical spur is smaller. The paramere’s shank of D. rusticus is also much more pilose. Neither species, Dysdercus chiriquinus nor D. rusticus, has a white annulus on the fourth antennal segment; both are different in color. Dysdercus stehliki sp.nov. is smaller than the other three species (Table 2), particularly so in D. longirostris and D. chiriquinus.

Table 2. Body lengths (in mm) of certain Dysdercus species with longer rostra. Note. The body widths of these species are, for Dysdercus longirostris, 4.5 mm (♂♂), and 6.0 mm (♀♀); for Dysdercus stehliki sp.nov., 4.1 mm (holotype ♂), and 5.2 mm (allotype ♀), measured at widest point of abdomen. The width measurements of D. longirostris were made by Dr. Luiz A. A. Costa (unpublished). The length measurements of the last three species were made by VAN DOESBURG (1968).

Males Females Dysdercus stehliki sp.nov. (all 10 types) 9.62–13.13 14.43–15.21 Dysdercus longirostris 13.0–15.0 15.0–19.0 Dysdercus chiriquinus 13.3–16.2 17.5–19.0 Dysdercus rusticus 12.0–15.0 13.0–18.0

Key to New World Dysdercus whose rostra extend at least to the fifth abdominal sternite 1 No white annulus on antennal segment IV...... 2. – With white annulus on antennal segment IV...... 3. 2 Black spot on each corium. ... Dysdercus chiriquinus Distant, 1883 – Without black spot on each corium...... Dysdercus rusticus Stål, 1870 3 Without scutellar depression; with very little pilosity on antennal segments III–IV...... Dysdercus longirostris Stål, 1861 – With scutellar depression; with abundant antennal pilosity...... Dysdercus stehliki sp.nov.

Similarities between Dysdercus stehliki sp.nov. and D. longirostris. I have received 3 female specimens of Dysdercus longirostris, identified by van Doesburg, from the Instituto Oswaldo Cruz (Rio de Janeiro) (see VAN DOESBURG 1968: 126). The three specimens were collected from two different places and in different years: two in 1935, in the State of Rio de Janeiro; and another in the State of Espirito Santo, collected in 1944. The two sites are about 350–400 km apart. Dysdercus longirostris is much closer in geography and phylogeny to D. stehliki sp.nov. than to D. chiriquinus or D. rusticus. Dysdercus longirostris, in the coastal areas

386 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Dysdercus stehliki sp.nov. of Brazil, “occupies only a very restricted area […] and its long rostrum suggests adaptation to some special host plant,” which may be Sterculia excelsa (VAN DOESBURG 1968: 180) – the same genus on which we have found D. stehliki sp.nov. Dysdercus stehliki sp.nov. is found beyond the coast of Brazil, and does not feed (as far as we know) on the same species of Sterculia. In both, the rostrum extends onto 6S in males and 7S in females, there is a white annulus on the fourth segment of the antenna, and there are no spots on the corium. FREEMAN’s (1947) account of the Old World species of Dysdercus, notes that in nearly all the species, the rostrum reaches onto 2S or occasionally 3S (22 of 25 species). In two Old World species, D. (Paradysdercus) sidae Montrouzier, 1861 and D. (Paradysdercus) longiceps Breddin, 1901, the rostrum reaches to 4S or sometimes 5S; in D. (Leptophthalmus) fuscomaculatus Stål, 1863 (= D. mesiostigma Distant, 1888), it reaches to 5S or 6S, apparently in both sexes (FREEMAN 1947). Thus the extension of the rostrum in all Dysdercus to 6S or 7S is unusual. Nevertheless, the origin of the New World species does not reflect this trend. Dysdercus originally was Old World, with subgenera in the Ethiopian and Oriental Regions. The Old World Dysdercus which entered the Neotropics (see STEHLÍK 1965a, b, and VAN DOESBURG 1968) originated in Ethiopian Region (the subgenus Dysdercus (Dysdercus)); but these Dysdercus do not have long rostra. A few Oriental Dysdercus do have long rostra (see above), but these are not the subgenera from which the New World Dysdercus originated. Because all of the Ethiopian Region Dysdercus have short rostra, the longer rostra must have arisen in the New World, and the occasional New World and the occasional Oriental longer rostra must have occurred separately. The white annulus on the fourth antennal segment occurs in all instars of Dysdercus stehliki sp.nov., from the first to the adult (unpublished). With one exception, Old World Dysdercus do not have this annulus: one African species has an annulus in the nymphal stages but not the adult (FREEMAN 1947). Van Doesburg suggests that this occurrence “may be a much older nymphal feature in the genus, becoming manifest in the adults in the New World” (VAN DOESBURG 1968: 173). It would be very interesting to discover if, in other Dysdercus, a nymphal white annulus occurs. In my borrowed specimens, the fourth antennal segments of Dysdercus longirostris are light to dark brown, as are the rest of the segments (sometimes a little more dark), but not “dark grey to blackish” as in the description of D. longirostris (VAN DOESBURG 1968: 124). This is true also of D. stehliki sp.nov. The “[u]pper surface of head, antennae, corners of pronotum, sterna, and sides of abdomen [are] minutely pilose” (VAN DOESBURG 1968: 126, of Dysdercus longirostris). This pilosity does not occur in D. stehliki sp.nov. and, moreover, I have not seen it in my specimens of D. longirostris. Differences between Dysdercus stehliki sp.nov. and D. longirostris. In all three specimens of Dysdercus longirostris the color is very much the same. In D. stehliki sp.nov. the dorsum of the head is red suffused broadly with black; the D. longirostris specimens are brownish red, but mostly brown. The predominant color of the prothoracic tergum and the corium is yellow-brown, but considerably paler than in Dysdercus stehliki

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Table 3. Differences between Dysdercus stehliki sp.nov. and D. longirostris Stål, 1861.

D. stehliki sp.nov. D. longirostris Color of prothoracic tergum pale yellow-brown darker, more brown and corium Head ventrally yellow-brown yellow Antennal segments III–IV more pubescence much less pubescence Antennal segment IV annulus white very pale yellow Scutellum some black markings no black markings Scutellar depression yes no Scent gland peritreme less rough than metasternum same roughness Explanate sternites more less Abdominal pits more developed less developed 6S & 7S abdominal pits circular shape obscured

sp.nov., which has more brown. The protergum’s posterior edge is white or pale yellow in both species; but just anterior to this margin is a light brown stripe (D. longirostris) or a black stripe (D. stehliki sp.nov.). The antennal segments of Dysdercus stehliki sp.nov. are very minutely pilose, especially the proximal half of the third and all of the fourth (Table 3); this pilosity is much less so in D. longirostris. The prothoracic disk of my Dysdercus longirostris specimens is not “livid to dirty yellow” (VAN DOESBURG 1968: 125), but light grayish yellow, like the corium. However, the scutellum of D. stehliki sp.nov. is darker (more orange) than the corium. In D. stehliki sp.nov., the scutellum has some black or grayish markings; these are absent in my borrowed specimens. On the abdomen’s sternites, laterally and often incorporating the spiracle (from 2S posteriorly), there is a small area, of the same color as the surrounding venter but of slightly different texture, slightly raised, and containing darker semi-circles surrounding it. These areas do not include the trichobothria, which are posterior to it, or the spiracles, which are anterior. These areas occur also in other Dysdercus as well. However, they differ in these two species (Table 3). In males [note: I had no specimens of male Dysdercus longirostris], the margins of 6S and 7S of Dysdercus stehliki sp.nov. are not enlarged (VAN DOESBURG 1968: 126) writes that they are enlarged in D. longirostris). The array of long conspicuous hairs on the ventral rims and the cuplike sclerites of D. stehliki sp.nov. is not mentioned by VAN DOESBURG (1968), although it does occur in other males of species of Dysdercus; in D. stehliki sp.nov. these hairs are much longer and more abundant than in other species of Dysdercus that I have seen. The paramere of D. stehliki sp.nov. has much fewer hairs,

388 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Dysdercus stehliki sp.nov. especially on its shank; and these hairs are smaller than in D. longirostris. VAN DOESBURG (1968) describes 2 small teeth on the head of the paramere of D. longirostris; in D. stehliki sp.nov. there may be 1–3 or even none (as in the holotype, Figs 3, 4). The spurs of the paramere of D. stehliki sp.nov. are also less sharply pointed (illustrations of D. longirostris by VAN DOESBURG (1968: figs. 177, 178) and another, unpublished, by Dr. Luiz A. A. Costa).

Notes on AHMAD & QADRI (2009). VAN DOESBURG (1968) listed four New World species that could not be grouped into his several species-categories: Dysdercus andreae (Linnaeus, 1758), D. ruficollis (Linnaeus, 1764), D. lunulatus Uhler, 1861, and D. mimuloides Blöte, 1933. AHMAD & QADRI (2009) have proposed a new subgenus – Dysdercus (Andreae) – for these three species (not including D. mimuloides), based exclusively on the species’ male and female genitalia. They do not formally name the subgenus, but they write, “D. andreae is hereby designated as its type species” (p. 129). Ahmad and Qadri also include D. mimulus Hussey, 1929, because they believe this species is an out-group of their new subgenus. Dysdercus mimuloides is mentioned in the Introduction to Ahmad and Qadri’s paper, but is not mentioned further (perhaps because they had no specimens to dissect?). VAN DOESBURG (1968) believes that D. andreae and D. mimuloides may be related, because the first is Caribbean and the second “occurs on the Pacific Coast near the border between Central and South America” (VAN DOESBURG 1968: 182) … . AHMAD & QADRI (2009) illustrate and describe the genitalia of these three species and D. mimulus; and I hope this new subgenus will promote further work on these four New World species, even though the original paper may be difficult to find. This is why I have included this discussion here.

Finally, and perhaps more important on Jaroslav Stehlík’s birthday, I think we should sing (perhaps with Brahms’ Opus 80): Quis confluxus hodie / Academicorum E longinquo convenerunt, / Protinusque successerunt In commune forum. Vivat nostra societas, / Vivant studiosi Crescat una veritas, / Floreat fraternitas Patriae prosperitas.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 389 C. W. SCHAEFER

Acknowledgements I am very grateful to Dr. Wagner d. S. Tavares (Universidade Federal de Viçosa, Minas Gerais, Brazil), who collected these Dysdercus in Minas Gerais; and to Dr. Petr Kment (Prague, Czech Republic), who helped me with this paper and others, and who shepherded me through this Festschrift. I am deeply grateful also to Dr. Luiz A. A. Costa (of the Museu Naçional da Universidade Federal do Rio de Janeiro), who sent to me some photographs and several drawings of Dysdercus longirostris, identified by van Doesburg; and to Dr. Hélcio Gil-Santana who answered questions about Brazilian geography and, with Dr. Cleber Galvão, shipped the specimens of D. longirostris to me (both of the Instituto Oswaldo Cruz, Rio de Janeiro). And finally I am indebted to my University of Connecticut colleagues, Ms V. Kask who drew the parameres, and Dr. J. O’Donnell who did wonderfully complicated things with photographs. Beyond all of that, I am deeply happy to dedicate this paper to Jaroslav L. Stehlík.

References

AHMAD I. & QADRI S. S. 2009. Male and female genitalial characters in the establishment of a new subgenus of cotton stainer’s genus Dysdercus Guérin-Meneville (Hemiptera: Pyrrhocoridae). Pakistan Journal of Zoology 41: 125–130. DOESBURG P. H. VAN 1968: A revision of the New World species of Dysdercus Guérin Méneville (Heteroptera, Pyrrhocoridae). Zoologische Verhandelingen 97: 1–215. FREEMAN P. 1947: A revision of the genus Dysdercus Boisduval (Hemiptera, Pyrrhocoridae), excluding the American species. Transactions of the Royal Entomological Society 98: 373–424. SCHAEFER C.W. 1977: Genital capsule of the trichophoran male (Hemiptera: Heteroptera: Geocorisae). International Journal of Insect Morphology and Embryology 6: 277–301. STEHLÍK J. L. 1965a: Mission Zoologique de l’I.R.S.A. en Afrique Orientale (P. Basilewsky—N. Leleup, 1957) Pyrrhocoridae (Het.). Acta Musei Moraviae, Scientiae Naturales 50: 211–250. STEHLÍK J. L. 1965b: Pyrrhocoridae and Largidae collected by E. S. Brown on Solomon Islands (Heteroptera). Acta Musei Moraviae, Scientiae Naturales 50: 253–292.

390 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 391–394, 2013

Spilostethus stehliki sp.nov. (Hemiptera: Heteroptera: ) from Ethiopia

JÜRGEN DECKERT Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt- Universität zu Berlin, Invalidenstraße 43, D-10115 Berlin; e-mail: [email protected]

DECKERT J. 2013: Spilostethus stehliki sp.nov. (Hemiptera: Heteroptera: Lygaeidae) from Ethiopia. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 391–394. – A new species of the genus Spilostethus Stål, 1868 (Hemiptera: Heteroptera: Lygaeidae: Lygaeinae), Spilostethus stehliki sp.nov., is described from the Simien Mountains in northern Ethiopia, and compared with Spilostethus taeniatus Stål, 1865, the species most similar to it.

Keywords. Heteroptera, , Lygaeidae, true bugs, taxonomy, new species, Afrotropical Region

Introduction Spilostethus Stål, 1868, is a group of aposematically coloured species, all from the Old World. At present, 24 species are known, including the new species described here (SLATER 1964a, SLATER & O’DONNELL 1995, LINNAVUORI 1978, CARAPEZZA 2002). A total of 17 species are now known from the Afrotropical Region. Three subspecies are described for S. rivularis (Germar, 1838) and two for S. furcula (Herrich-Schaeffer, 1850); see also LINNAVOURI (1974, 1978). The species are known for their striking similarities in coloration and mimicry. Nothing has emerged to challenge what SLATER (1964b) said of them half a century ago: “A number of interesting problems of speciation, variation, mimicry, sex recognition, etc., appear to await the careful investigator.” The main sources of information for the African Spilostethus are the publications by SLATER (1964a, b) and SLATER & SPERRY (1973), both focused on southern Africa, and by LINNAVUORI (1974, 1978). In this paper one additional species is described from mountains of northern Ethiopia.

Taxonomy Spilostethus stehliki sp.nov. (Figs 1, 2, 5) Type material. Holotype: Male. ETHIOPIA: ‘ÄTHIOPIEN 15 Jan. 1996, Simien Mountains, östl. Debark Sankaber Camp, 13°13′ N / 28°02′E, leg. J. Deckert, 3245 m’. Paratypes: 4 females, same locality and date. The type material is deposited in the Museum für Naturkunde, Berlin. Description. Coloration. General coloration black and orange-red. Head with Y-shaped black pattern; tylus and vertex black, except for base orange-red dorsally; juga and lateral and ventral parts of antenniferous tubercles orange-red. Antennae, rostrum, and legs unicolourous black, coxae partly paler. Pronotum black with three narrow, longitudinal,

391 J. DECKERT orange-red stripes (one median orange-red stripe, prolonged to basal midway of the black scutellum, and two stripes on lateral margins); corium dark except for orange-red stripe at the side and on the inner part at the level of claval commissure; clavus black, borders of scutellum orange-red, membrane pale grey to white, except for black area adjacent to corium. Pleura of each thoracic segment with orange-red patches, abdominal segments black with orange-red regions. Structure. Head convex, moderately longer than wide, rostrum ends between middle and hind coxae. Pronotum without prominent longitudinal carinae, lateral margin somewhat swollen, peritreme of metathoracic scent gland formed by shallow groove, without auricle. Scutellum with ridge at the middle part, especially at apex. Femora and tibiae of male with small but distinct spines on inner surface. Submacropterous, wings not reaching apex of abdomen, membrane slightly reduced. Pilosity. Body covered with short, semi-erect, silvery hairs. Sculpture. Anterior and posterior area of pronotum with a few black punctures, also some black punctures on first pleura. Shape of male parameres as in Fig. 5. Measurements (mm). Male (holotype). Body length 7.0; head: width (including eyes) 1.5; interocular width 1.0; length of antennomeres: 1 – 0.5, 2 – 1.0, 3 – 0.7, 4 – 1.1; pronotum: length at centre 1.5, width across humeral edges 2.2; scutellum: length 0.8, width 1.1; corium: length 1.7. Females (n = 4). Body length 6.9 (6.7–7.4); head: width (including eyes) 1.5; interocular width 1.0; length of antennomeres: 1 – 0.4 (0.39–0.42), 2 – 0.98 (0.93–1.0), 3 – 0.7 (0.6–0.75), 4 – 1.02 (1.0–1.05); pronotum: length at centre 1.35 (1.1–1.42), width across humeral edges 2.2 (2.0–2.36); scutellum: length 0.89 (0.86–0.92), width 1.2 (1.0–1.27); corium: length 1.8 (1.6–2.0). Differential diagnosis. Spilostethus stehliki sp.nov. differs from other species of Spilostethus in its slightly reduced wings in combination with the colour pattern, smaller size (of about 7 mm), and the shape of parameres. Etymology. The species is dedicated to Jaroslav L. Stehlík for the occasion of his 90th birthday. Biology. The species was collected on the ground in a dry meadow and was found together with two other lygaeine bugs, Gondarius inexpectatus Štys, 1972 and Spilostethus amaenus (Bolívar, 1897). Distribution. The species is known from only its type locality in northern Ethiopia, the Simien Mountains, at an altitude of over 3000 m.

Discussion SLATER (1964b) mentioned that Spilostethus taeniatus Stål, 1865, is “most closely related to S. trilineatus (Fabricius, 1794) from which species it may however be readily distinguished by the length of the median red pronotal stripe that attains the anterior margin of the pronotum […]”. LINNAVUORI (1974) discussed and keyed the red-coloured species of the African rivularis species-group of Spilostethus with the species S. rivularis (Germar, 1838), S. taeniatus (Stål, 1865), S. trilineatus (Fabricius, 1794), and S. crudelis

392 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Spilostethus stehliki sp.nov.

Figs 1–6. Spilostethus spp. 1, 2, 5 – S. stehliki sp.nov., holotype: 1– dorsal view; 2 – lateral view; 5 – paramere. 3, 6 – S. taeniatus (Stål, 1865): 3 – holotype (Naturhistoriska riksmuseet, Stockholm), dorsal view; 6 – specimen from Mozambique: Delagoa Bay (Naturhistorisches Museum Wien), paramere. 4 – S. rivularis epimetheus Linnavuori, 1974, holotype (American Museum of Natural History, New York), dorsal view. Scale bars: 1 mm (1–4), 0.1 mm (5, 6).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 393 J. DECKERT

(Fabricius, 1781). He keyed Spilostethus taeniatus (but did not illustrate it) as follows: “Pronotum with a percurrent narrow red median stripe. Scutellum also with a red midline”, giving the distribution as South Africa, Sudan, and Ethiopia. Further, LINNAVUORI (1978) mentioned S. taeniatus for South Sudan. The species appears to be widely distributed in southern and tropical Africa; however, identification based on colour patterns alone has to be further validated. Spilostethus stehliki sp.nov. from the Simien Mountains has generally a similar colour pattern to that found in the S. rivularis species-group. Comparison with the type material of S. taeniatus (Fig. 3), held in the Naturhistoriska riksmuseet in Stockholm, and with redescriptions by SLATER (1964) makes it clear that S. taeniatus and other similar known species are quite different from Spilostethus stehliki sp.nov. At first sight Spilostethus stehliki sp.nov. resembles S. taeniatus, but in S. stehliki the dark coloration on the pronotum, corium and ventral parts is predominant and the black corial stripes reach the membrane. In S. taeniatus the orange-red areas of the body are about equal to the black parts (Fig. 3). The newly-described species also differs in the shape of the parameres (Figs 5–6) and size (7–9 mm). Other species of the S. rivularis species group are S. rivularis (with subspecies rivularis epimethaeus Linnavouri, 1974, see Fig. 4), S. lemniscatus (Stål, 1855) and S. trilineatus, but all of them have parameres sufficiently distinctive for clear discrimination.

Acknowledgements First of all, I must thank Jaroslav L. Stehlík for his numerous essential and stimulating publications on Pyrrhocoridae, Largidae and other groups. For the opportunity to study museum collections, I thank Herbert Zettel (Vienna), Randall Toby Schuh (New York), and Bert Viklund (Stockholm). Petr Kment (Praha) and Elöd Kondorosy (Keszthely) kindly reviewed the text and provided useful improvements to the manuscript.

References

CARAPEZZA A. 2002: Heteroptera of Jordan: New Taxa and new records (Hemiptera Heteroptera). Naturalista Siciliano, Seria IV 26: 35–76. LINNAVUORI R. 1974: Studies on Palearctic and African Heteroptera. Acta Zoologica Fennica 30: 1–35. LINNAVUORI R. 1978: Hemiptera of the Sudan, with remarks on some species of the adjacent countries 6. Acta Zoologica Fennica 153: 1–108. SLATER J. A. 1964a: A catalogue of the Lygaeidae of the World, I. Baltimore, Maryland, 778 pp. SLATER J. A. 1964b: Hemiptera (Heteroptera) Lygaeidae. Pp. 15–228. In: HANDSTRÖM B., BRINCK P. & RUDEBECK G. (eds.): South African animal life. Results of the expedition in 1950–1951. Vol. 10, Swedish Natural Science Council, Stockholm, 228 pp. SLATER J. A. & SPERRY B. 1973: The biology and distribution of the South African Lygaeinae, with descriptions of new species (Hemiptera, Lygaeidae). Annals of the Transvaal Museum 28: 117–204. SLATER J. A. & O’DONNELL J. E. 1995: A Catalogue of the Lygaeidae of the World (1960-1994). New York Entomological Society, New York, 410 pp.

394 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 395–406, 2013

Description of four new species of Ninyas from Venezuela, a key to the known species and some new records (Hemiptera: Heteroptera: )

HARRY BRAILOVSKY Departamento de Zoología, Instituto de Biología, UNAM, Apdo Postal 70-153, México, D.F. México; e-mail: @ibiologia.unam.mx

BRAILOVSKY H. 2013: Description of four new species of Ninyas from Venezuela, a key to the known species and some new records (Hemiptera: Heteroptera: Geocoridae). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 395–406. – Four new species of Ninyas Distant, 1882 (Geocoridae), N. nigricans sp.nov., N. reclusus sp.nov., N. solitarius sp.nov. and N. stehliki sp.nov., all from Venezuela, are described. New distributional records for N. distanti Montandon, 1907 from Peru and N. torvus Distant, 1893 from Venezuela are added. A key to the 14 known species of Ninyas is included. Sexually dimorphic features are commented upon. Keywords. Insecta, Hemiptera, Heteroptera, , Geocoridae, Ninyas, new species, new records, key, dimorphism, Venezuela, Neotropical Region

Introduction The genus Ninyas Distant, 1882 is currently classified in the family Geocoridae, superfamily Lygaeoidea (HENRY 1997). Members of this genus are small to medium- sized (3–5.5 mm) and are recognized by: “polished” body; eyes pedunculate, reniform, transversely straight, and remote from anterior angles of pronotum; portion of head bearing eyes extending laterally and constricted; ocellus closer to medial line; claval commissure well defined; outer and inner margins and apical third of clavus punctate; corium with a row of punctures along margin adjacent to clavus and another row of punctures extending from base approximately through central area and curving towards apex of corium. All the species of Ninyas are similar in shape and are variable in colour, and in addition to the variation in colour there is a striking difference in some populations in the degree of opacity of the corium. Most specimens have a transparent corium but in some the corium is completely opaque. Members of this genus are usually taken by beating from trees and shrubs (BARANOWSKI & SLATER 2005). Ninyas currently contains 10 described species widely distributed in the Neotropical region including the West Indies (Great and Lesser Antilles): N. deficiens (Lethierry, 1881) (Bahama Islands, Cuba, Dominica, Dominican Republic, Grenada, Guadeloupe Island, Haiti, Jamaica, Puerto Rico, and St. Lucia); N. distanti Montandon, 1907 (Brazil); N. dominicanus Baranowski, 2005 (Dominican Republic); N. humeralis Barber, 1947 (Cuba); N. obrieni Baranowski, 2005 (Puerto Rico); N. punctatus Baranowski, 2005 (Martinique); N. solubilis Distant, 1893 (Panama); N. strabo Distant, 1882 (Cuba, Dominica, Grenada, Hispaniola, Jamaica, Mexico, Panama, and Puerto Rico); N. torvus

395 H. BRAILOVSKY

Distant, 1893 (Panama); and N. woodruffi Baranowski, 2005 (Dominican Republic) (BARANOWSKI & SLATER 2005; BARBER 1939, 1947; BARBER & ASHLOCK 1960; DISTANT 1882, 1893; FROESCHNER 1999; MONTANDON 1907; SLATER 1964). Ninyas has not been previously recorded from Venezuela. In the present paper four new species are described and in addition N. torvus is here cited for the first time from Venezuela, and N. distanti from Peru. A key to all known species is included and comments on sexual dimorphism are added.

Materials and methods The material examined is deposited in the following collections: AMNH ...... American Museum of Natural History, New York, USA BMNH ...... Natural History Museum, London, United Kingdom HNHM ...... Hungarian Natural History Museum, Budapest, Hungary IZAV ...... Instituto de Zoología Agrícola, Universidad Central de Venezuela, Maracay, Edo. Aragua, Venezuela TAMU ...... Texas A & M University (College Station), Texas, USA UCDC ...... University of California, Davis, Bohart Museum of Entomology, California, USA UNAM ...... Instituto de Biología, Universidad Nacional Autónoma de México, México City, México USNM ...... Smithsonian Institution, National Museum of Natural History, Washington, D.C., USA

Taxonomy Ninyas stehliki sp.nov. (Fig. 10)

Type material. HOLOTYPE: ♂, Venezuela: Estado de Aragua: Parque Nacional Rancho Grande, Estación de Biología, 1,100 m a.s.l., 16.vi.1983, C. Bordon (UNAM). PARATYPES: Venezuela: Aragua: Rancho Grande, 20 km NW of Maracay, 1,100 m a.s.l., at lights, 23.vii.1976, 1 ♂, M. H. Sweet (TAMU); Rancho Grande, 1,300 m a.s.l., 14.vii.1979, 1 ♂, R. O. Schuster and A. Grigarick (UCDC); Rancho Grande, 1,100 m a.s.l., 20.ix.1949, 1 ♂ 1 ♀, F. Fernandez and P. Fenives (IZAV); Rancho Grande, 1,100 m a.s.l., 1.iv.1966, 1 ♂ 1 ♀, J. and B. Bechyne (IZAV); Rancho Grande, 1,100 m a.s.l., 7.vii.1976, 1 ♀, J. Gonzalez (IZAV); Rancho Grande, 1200–1400 m a.s.l., 13.viii.1988, 4 ♀♀, C. W. and L. O’Brien and G. J. Wibmer (UNAM); Villa de Gura, Est. Exp. Cataurito, 1.000 m a.s.l., 19.vi.1985, 2 ♂♂ 1 ♀, 16.i.1980, 1 ♀, J. L. Garcia (IZAV); Portachuelo, 1,120 m a.s.l., 14.viii.1988, 2 ♀♀, C. W. and L. O’Brien and G. J. Wibmer (UNAM); El Limon, 850 m a.s.l., 23.ix.1970, 1 ♀, J. and B. Bechyne (IZAV). Carabobo: Canoabo, 19.iii.1969, 1 ♂ 1 ♀, J. and B. Bechyne (IZAV); San Esteban (Las Quiguas), 184 m a.s.l., 5.–13.xi.1974, 1 ♀, J. Salcedo, R. Dietz and J. L. Garcia (IZAV). Description. A relatively large species, up to 5 mm; elongate, parallel-sided, lateral corial margins weakly emarginate. Male. Dorsal coloration. Head including the space behind eyes completely light pale yellow; eyes dull reddish; ocellar tubercle dull to light reddish; antennal segment I light pale yellow, segment II light brown, III with anterior half light brown and posterior half light pale yellow, and IV light brown with basal third light pale yellow. Anterior and posterior lobes of pronotum light pale yellow; darkened area restricted to a series of

396 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Ninyas punctures behind collar and a second transverse series behind callar area; posterior lobe with irregular, coarse, castaneous punctures; humeral angles with a light brown macula. Scutellum castaneous, heavily punctate except for an elevated, Y-shaped, yellowish- white callose area occupying much of scutellar disk. Clavus castaneous, opaque; corium transparent with a faint castaneous tinge; apical margin of corium with an incomplete brown band; hemelytral membrane transparent with a faint castaneous tinge, and with basal angle dark brown. Connexival segments and abdominal terga light pale yellow. Ventral coloration. Including rostral segments (apex of IV light reddish), and legs (last tarsal segment light brown) light pale yellow; pro-, meso-, and metapleura with castaneous punctures; middle third of abdominal sterna with a faint castaneous tinge. Structure. Head gently downward-sloping, slightly more than two times wider across eyes than long; clypeus extending just beyond middle of antennal segment I; rostrum reaching posterior third of mesosternum or anterior third of metasternum. Pronotum trapezoidal, bilobate; lateral margin of anterior lobe convex; lateral margins of posterior lobe straight; humeral angles obtuse; posterior margin almost straight. Hemelytral membrane extending markedly beyond apex of last abdominal segment. Sculpture. Dorsal surface glabrous; head ventrally glabrous; thorax with a few erect hairs; femora moderately pilose with short, semi-erect setae; tibiae with long, erect setae; middle third of abdominal sternite III to VII covered with long, erect, dense and heavily compact setae. Head and abdomen impunctate. Thoracic pleura coarsely punctate. Female. Coloration. Similar to male holotype. Dorsal abdominal segments and genital plates light pale yellow with faint castaneous tinge. Sculpture similar to male holotype except the middle third of abdominal sternite III to VII with short, scattered, semi-erect setae. Variation. i) Antennal segment II light pale yellow and IV yellow with apical third light brown. ii) Rostral segments I–II light pale yellow, and III–IV light brown. iii) Acetabulae yellowish white. iv) Mesosternum castaneous. v) Pro-, meso-, and metapleura with reddish-brown punctures. Measurements (mm). One male first, then one female. Body length 5.28, 5.30; head: length 0.72, 0.78, width (including eyes) 1.56, 1.62, preocular distance 0.40, 0.45, interocellar distance 0.44, 0.47; length of antennal segments: I – 0.32, 0.32, II – 0.64, 0.68, III – 0.60, 0.60, IV – 0.76, 0.64; pronotum: maximum length of anterior lobe 0.32, 0.29, maximum length of posterior lobe 0.64, 0.72, maximum width of anterior lobe 1.20, 1.20, maximum width of posterior lobe 1.56, 1.68; scutellum: length 0.72, 0.76, width 0.76, 0.84; length of claval commissure 0.32, 0.36. Differential diagnosis. Ninyas distanti (Fig. 2) known from Brazil and N. torvus (Fig. 12) from Panama are species longer than 4.5 mm, sharing with N. stehliki sp.nov. the scutellum bearing an elevated and complete Y-shaped, yellowish-white callose area occupying much of scutellar disk, and the hemelytral membrane conspicuously extending beyond the apex of last abdominal segment and without a dark stripe running through the middle. In N. stehliki sp.nov. (Fig. 10) described from Venezuela, the dorsal surface of head including the space behind eyes, and the anterior and posterior lobe of pronotum are light

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 397 H. BRAILOVSKY pale yellow, the basal angle of the hemelytral membrane is dark brown, and the humeral angles bear a light brown macula. In N. distanti the dorsal surface of head is dark brownish with apical portion pale light yellow, the anterior lobe of pronotal disk is entirely dark brownish-black, the posterior lobe bears a quadrate dark brown spot, the basal angle of the hemelytral membrane is castaneous-transparent, and the humeral angles are yellow without light brown macula. In N. torvus the dorsal surface of head is pale reddish, the anterior lobe of the pronotum is pale reddish and laterally pale yellow, the middle third of the posterior lobe bears a quadrate, pale reddish spot, the basal angle of the hemelytral membrane is castaneous-transparent, and the humeral angles are yellow without light brown macula. In addition, the dorsal abdominal segments of N. stehliki sp.nov. are entirely light pale yellow, while in the other two species those are yellow with black marks. Etymology. I take this opportunity to name this species in honour of Dr. Jaroslav L. Stehlík, the distinguished Czech hemipterist, in recognition of his great entomological work, particularly on Largidae and Pyrrhocoridae. Distribution. Known only from Venezuela (Aragua and Carabobo).

Ninyas nigricans sp.nov. (Fig. 4)

Type material. Holotype: ♀, Venezuela: Carabobo: Güique, 23.ii.1968, J. and B. Bechyne (IZAV). Description. Small species, less than 3.50 mm; elongate, ovoid, lateral corial margins distinctly emarginate. Female. Dorsal coloration. Head including the space behind eyes completely light pale yellowish-castaneous; eyes and ocellar tubercle light reddish; antennal segments I–III pale yellowish castaneous and IV pale yellowish castaneous with apical third pale brown. Anterior and posterior lobe of pronotum light pale yellowish-castaneous; darkened area restricted to a series of punctures behind collar and a second transverse series behind callar area; posterior lobe with irregular, coarse, dark castaneous punctures; humeral angles with a light brown macula. Scutellum dark reddish-brown with apex pale yellowish-castaneous; impunctate except lateral margins with a row of punctures. Clavus and corium pale yellowish-castaneous; apical margin of corium with an incomplete brown band; hemelytral membrane transparent with a faint castaneous tinge. Connexival segments pale yellowish-castaneous; terga pale yellowish-castaneous with irregular dark marks. Ventral coloration. Including rostral segments and legs pale yellowish-castaneous; pro-, meso-, and metapleura with dark castaneous punctures; mesosternum and genital plates dark reddish brown. Structure. Head gently downward-sloping, broad, more than two times as wide across eyes than long; clypeus reaching apex of antennal segment I; rostrum reaching anterior third of metasternum. Pronotum trapezoidal, bilobate; lateral margin of anterior lobe convex; lateral margins of posterior lobe straight; humeral angles obtuse; posterior

398 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Ninyas margin straight. Hemelytral membrane slightly extending beyond apex of last abdominal segment. Sculpture. Dorsal surface glabrous; head ventrally glabrous; thorax almost glabrous; femora moderately pilose with short, semi-erect setae; tibiae with long, erect setae; abdominal sternites III to VII covered with short to medium-sized, scattered, semi-erect setae. Head and abdomen impunctate. Thoracic pleura coarsely punctate. Male. Unknown. Measurements (mm). Body length 3.35; head: length 0.48, width (including eyes) 1.24, preocular distance 0.30, interocellar distance 0.20; length of antennal segments: I – 0.16, II – 0.40, III – 0.36, IV – 0.48; pronotum: maximum length of anterior lobe 0.24, maximum length of posterior lobe 0.48, maximum width of anterior lobe 0.80, maximum width of posterior lobe 1.20; scutellum: length 0.52, width 0.56; length of claval commissure 0.19. Differential diagnosis. Ninyas nigricans sp.nov. (Fig. 4) is different from all other known species of Ninyas in having the scutellar disk dark reddish-brown with apex pale yellowish-castaneous and lacking a Y-shaped callose area. In the other known taxa the scutellum is pale yellow with complete or incomplete Y-shaped yellowish-white callose area occupying much of scutellar disk. Etymology. Latin participle nigricans (= blackened). The specific name refers to the dark reddish-brown scutellum. Distribution. Known only from Venezuela (Carabobo).

Ninyas reclusus sp.nov. (Fig. 7)

Type material. Holotype: ♂, Venezuela: Aragua: Choroni, 1,600 m a.s.l., 2.ii.1970, J. and B. Bechyne (IZAV). Description. A relatively large species, up to 5 mm; elongate, parallel-sided, lateral corial margin emarginate. Male. Dorsal coloration. Vertex and posterior portion of head, including the space behind eyes, lustrous black; frons and clypeus lustrous orange; eyes and ocellus pale reddish; antennal segment I pale yellowish-orange with apical border pale reddish, segment II dark castaneous with apical third brown, III with anterior half brown and posterior half castaneous orange, and IV dark brown with basal third castaneous orange. Anterior lobe of pronotum lustrous brown with frontal angles and a wide median transverse band, pale yellow; posterior lobe pale yellow with punctures, and two quadrate spots lateral to middle line lustrous brown; humeral angles with pale lustrous brown macula. Scutellum dark reddish-brown, heavily punctate, except for an elevated, Y- shaped, yellowish-white callose area occupying much of scutellar disk. Clavus dark lustrous brown with basal third pale yellow; corium opaque, pale yellowish-white with dark brown punctures; apical margin of corium with an incomplete dark brown band; hemelytral membrane transparent with a faint castaneous tinge. Connexival segments pale yellow; abdominal terga dark brown with middle third of segments V–VI, and lateral margins of segments VI–VII pale yellow.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 399 H. BRAILOVSKY

Ventral coloration. Dark lustrous blackish-brown with following areas pale yellow: rostral segments I–IV (apex of IV brown), lateral area of head including the buccula, collar, anterior and posterior border of prothorax, metathoracic peritreme, all acetabula, legs, posterior angle of metathorax, and pleural margin of abdominal sterna III–VII. Structure. Head gently downward-sloping, slightly more than two times wider across eyes than long; clypeus extending just beyond mid-point of antennal segment I; rostrum reaching posterior third of metasternum. Pronotum trapezoidal, bilobate; lateral margin of anterior lobe convex; lateral margins of posterior lobe straight; humeral angles obtuse, posterior margin almost straight. Hemelytral membrane conspicuously extending beyond apex of last abdominal segment. Sculpture. Dorsal surface glabrous; head ventrally and thorax glabrous; femora and tibiae with long, erect setae; middle third of abdominal sternites III to VII covered with long, erect, dense and heavily compact setae. Head and abdomen impunctate. Thoracic pleura coarsely punctate. Measurements (mm). Body length 5.20; head: length 0.76, width (including eyes) 1.64, preocular distance 0.38, interocellar distance 0.29; length of antennal segments: I – 0.28, II – 0.68, III – 0.56, IV – 0.76; pronotum: maximum length of anterior lobe 0.34, maximum length of posterior lobe 0.62, maximum width of anterior lobe 1.20, maximum width of posterior lobe 1.56; scutellum: length 0.74, width 0.76; length of claval commissure 0.27. Female. Unknown. Differential diagnosis. Ninyas reclusus sp.nov. is related to N. dominicanus, the two sharing the vertex and posterior portion of head including the space behind eyes lustrous black, and frons and clypeus lustrous orange to yellow, the humeral angles with a pale brown macula, and the hemelytral membrane without median dark stripe. Ninyas reclusus sp.nov. (Fig. 7) is longer than 5 mm, and distinguished by having the posterior margin of pronotum yellow with humeral angles and middle third brown, the middle third of pronotal disk with a yellow longitudinal band, and the clavus dark lustrous brown with the basal third pale yellow. N. dominicanus, described from the Dominican Republic, is shorter than 4.10 mm, the posterior margin of pronotum has a brown transverse band, the middle third of the pronotal disk features a brown longitudinal band, and the clavus is dark brown. Etymology. Latin adjective reclusus, -a, -um; the specific name refers to the hidden nature of this species. Distribution. Known only from Venezuela (Aragua).

Ninyas solitarius sp.nov. (Fig. 8)

Type material. Holotype: ♂, Venezuela: Territorio Federal Amazonas: Cerro de la Neblina, Camp X, 1690 m a.s.l., 0°54′N-60°2′W, 12.ii.1985, W. E. Steiner (USNM). Description. Small species, less than 3.50 mm long; more or less quadrate, lateral corial margins distinctly emarginate.

400 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Ninyas

Male. Dorsal coloration. Head including the space behind eyes completely light pale yellow; eyes dull reddish; antennal segments I–II pale yellow, segment III–IV pale castaneous orange. Anterior and posterior lobes of pronotum light pale yellowish-white; darkened area restricted to a series of castaneous punctures behind collar and a second transverse series rearwards of callar area; posterior lobe with irregular, coarse, castaneous punctures; humeral angles with brown macula. Scutellar disk with irregular, coarse, dark castaneous-to-brown punctures; middle line with wide, longitudinal, yellowish-white callose area, and basally with the lateral angles dark brown. Clavus yellowish-white, opaque; corium yellowish-white, opaque, with endocorium mostly transparent; apical margin of corium yellowish white, with only the apical angle brown; hemelytral membrane transparent with a faint castaneous tinge. Connexival segments pale yellowish-castaneous; abdominal terga dark brown, except terga V–VI mostly yellowish. Ventral coloration. Head, rostral segments (apex of IV pale brown), collar, acetabula, coxae, trochanter, tibiae, tarsi, posterior margin of metapleura, and metathoracic scent gland peritreme, pale yellowish-white; femora pale orange with basal third pale yellowish-white; mesosternum blackish-brown; prosternum, metasternum, propleura and mesopleura yellowish white with punctures dark castaneous; metapleura dark castaneous orange; evaporatorium dark brown; abdominal sterna III–VII dark orange with pleural margin yellowish-white; genital capsule dark orange. Structure. Head almost flat, not downward-sloping, more than two times as wide across eyes than long; clypeus extending just beyond mid-point of antennal segment I; rostrum reaching posterior third of metasternum. Pronotum trapezoidal, bilobate; lateral margin of anterior lobe convex; lateral margins of posterior lobe straight; humeral angles obtuse; posterior margin almost straight. Hemelytral membrane extending markedly beyond apex of last abdominal segment. Sculpture. Dorsal surface glabrous; head ventrally and thorax glabrous; femora and tibiae pilose with long, erect setae; middle third of abdominal sternites III to VII covered with long, erect, dense and heavily compact setae. Head and abdomen impunctate. Thoracic pleura coarsely punctate. Measurements (mm). Body length 3.26; head: length 0.52, width (including eyes) 1.32, preocular distance 0.32, interocellar distance 0.24; length of antennal segments: I – 0.22, II – 0.36, III – 0.36, IV – 0.56; pronotum: maximum length of anterior lobe 0.24, maximum length of posterior lobe 0.48, maximum width of anterior lobe 0.96, maximum width of posterior lobe 1.24; scutellum: length 0.52, width 0.56; length of claval commissure 0.18. Female. Unknown. Differential diagnosis. Ninyas solitarius sp.nov. (Fig. 8) is related to N. deficiens (Fig. 1). Both species have the head dorsally, including the space behind eyes, completely pale yellow, the humeral angles with brown macula, the hemelytral membrane without dark median irregular stripe, the scutellar disk with wide, longitudinal, yellowish-white callose area, and both are small species, measuring less than 3.60 mm. In N. solitarius sp.nov. the clavus is yellowish-white, opaque, the corium yellowish- white, opaque, except for endocorium transparent, and apical margin of corium

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 401 H. BRAILOVSKY yellowish-white, opaque, with only the apical angle brown. In N. deficiens, recorded from West Indies, the clavus is dark castaneous, the corium transparent, and the apical margin of the corium has an incomplete brown band. Ninyas strabo (Fig. 11) is also related to the new species but the space behind its eyes is dark brown. Etymology. Latin adjective solitarius, -a, -um (= solitary), as this species is known only from a single male. Distribution. Known only from Venezuela (Territorio Federal del Amazonas).

Ninyas distanti Montandon, 1907 (Fig. 2) Ninyas distanti Montandon, 1907: 293–295.

Type material examined. Holotype ♂: Brazil: Goyas: Itahy (HNHM). New records. Peru: Junin: San Ramón de Pangoa, 40 km SE Satipo, 750 m a.s.l., 7.iii.1972, 1 ♂, R. T. & J. C. Schuh (AMNH); Satipo, xi.1942, 1 ♂, Paprzycki (UNAM). Diagnosis. A large species, up to 5 mm, parallel-sided, characterized by head dorsally light pale yellow with vertex and posterior portion entirely lustrous black; scutellum black with a complete, Y-shaped, yellowish-white callose area occupying much of scutellar disk; pronotal lobe lustrous black to brown with lateral margins yellow; humeral angles without black or dark brown spot; and apical angle of corium with wide brown area. Distribution. Recorded from Brazil (MONTANDON 1907) and Peru.

Ninyas torvus Distant, 1893 (Fig. 12) Ninyas torvus Distant, 1893: 388–389.

Type material examined. Paratype: Panamá: Bugaba, 900–1500 ft, 1 ♂, Champion lgt. (BMNH). New records. Venezuela: Distrito Federal: Chichiriviche, Petaquire, 27.i.1977, 1 ♂, 1 ♀, C. J. Rosales and L. J. Joly (IZAV, UNAM). Diagnosis. A large species, up to 4.5 mm, parallel-sided, with lateral corial margin emarginate and characterized by hemelytral membrane without dark stripe running through the middle; humeral angles of pronotum with brown macula; head dorsally including the space behind eyes pale reddish; pronotum pale yellow, with punctures reddish-brown to pale castaneous, and with middle third of anterior lobe and a quadrate and wide longitudinal band running across the middle third pale reddish; scutellum with a complete, Y-shaped, yellowish-white callose area occupying much of scutellar disk; and apical corial margin with incomplete brown band. Distribution. Previously known from Panama (Volcan de Chiriqui, David and Bugaba) (DISTANT 1893). It is recorded here for the first time from Venezuela.

402 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Ninyas

Figs. 1–13. Habitus of Ninyas spp. 1 – N. deficiens (Lethierry, 1881), ♂; 2 – N. distanti Montandon, 1907, ♂; 3– N. humeralis (Barber, 1947), ♀; 4 – N. nigricans sp.nov., ♀; 5 – N. obrieni Baranowski, 2005, ♀; 6 – N. punctatus Baranowski, 2005, ♂; 7 – N. reclusus sp.nov., ♂; 8 – N. solitarius sp.nov., ♂; 9 – N. solubilis Distant, 1893, ♀; 10 – N. stehliki sp.nov., ♀; 11 – N. strabo Distant. 1882, ♀; 12 – N. torvus Distant, 1893, ♂; 13 – N. woodruffi Baranowski, 2005, ♂.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 403 H. BRAILOVSKY

Key to the species of Ninyas Distant, 1882 Modified from BARANOWSKI & SLATER (2005): 1 Hemelytral membrane with a distinct dark median longitudinal stripe...... 2 – Hemelytral membrane sometimes with infuscated areas, but never with a complete dark stripe running the entire length of the membrane. ... 3 2 Humeral angles of pronotum with a red macula; head dorsally, including the space behind eyes, completely pale yellow (Fig. 3)...... N. humeralis (Barber, 1947) – Humeral angles of pronotum lacking red macula; head dorsally pale yellow with vertex and posterior portion lustrous black (Fig. 13)...... N. woodruffi Baranowski, 2005 3 Head pale reddish; anterior lobe of pronotum almost entirely pale reddish (Fig. 12)...... N. torvus Distant, 1893 – Head completely pale yellow or pale yellow with vertex and posterior portion black; anterior lobe of pronotal disk pale yellow with or without black transverse band or mostly black...... 4 4 Head dorsally pale yellow with the space behind eyes or interocular part entirely black...... 5 – Head dorsally including the space behind eyes completely pale yellow...... 9 5 Head dorsally pale yellow except the space behind eyes black...... 6 – Head dorsally black with apical third including the clypeus pale yellow...... 7 6 Abdominal sterna III to VII pale yellow; antennal segment IV yellow with apical third pale brown (Fig. 11)...... N. strabo Distant, 1882 – Abdominal sterna III to VII dark brown with pleural margin pale yellow; antennal segment IV pale brown (Fig. 9)...... N. solubilis Distant, 1893 7 Scutellum black with a median yellowish-white vitta that is not expanded laterally toward apical third...... N. dominicanus Baranowski, 2005 – Scutellum black with an elevated, Y-shaped, callose yellowish-white area...... 8 8 Humeral angles of pronotum without black spot; antennal segment IV yellow; anterior lobe of pronotal disk brown; posterior pronotal lobe brown with lateral margins yellow (Fig. 2)...... N. distanti Montandon, 1907 – Humeral angles of pronotum with brown spot; antennal segment IV dark brown with basal third castaneous orange; anterior lobe of pronotal disk brown with frontal angles and a median, transverse, wide,

404 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Ninyas

band pale yellow; posterior pronotal disk yellow with punctures and two quadrate spots lateral to middle line brown (Fig. 7)...... N. reclusus sp.nov. 9 Humeral angles of pronotum without brown macula (Fig. 5); apical corial margin without dark coloration. ... N. obrieni Baranowski, 2005 – Humeral angles of pronotum with brown spot; apical corial margin partly brown...... 10 10 Scutellum dark reddish brown with apex yellowish-castaneous (Fig. 4)...... N. nigricans sp.nov. – Scutellum pale to dark brown with a complete or incomplete Y-shaped, yellowish-white callose area occupying much of the scutellar disk...... 11 11 Brown band on apical corial margin complete (Fig. 6)...... N. punctatus Baranowski, 2005 – Brown band on apical corial margin incomplete...... 12 12 Total length of body longer than 5 mm (Fig. 10)...... N. stehliki sp.nov. – Total length of body less than 4 mm...... 13 13 Apical margin of corium with an incomplete brown band; clavus dark to pale castaneous (Fig. 1)...... N. deficiens (Lethierry, 1881) – Apical margin of corium yellowish-white opaque, with only the apical angle brown; clavus yellowish-white opaque (Fig. 8)...... N. solitarius sp.nov.

Sexual dimorphism Sexual dimorphism has not previously been reported in the genus Ninyas. During this revision, in which 13 of 14 known species were examined, it was noticed that in males the middle third of abdominal sternites III to VII is covered with a dense pilosity of long, silvery, erect and densely adpressed hairs; whereas on females the pilosity in that area consists of short hairs, sparse and semi-decumbent to completely decumbent. This dimorphism in pilosity might indicate a possible sensory function during the courtship and mating or it could be related to substrate recognition.

Acknowledgements I wish to thank the following individuals and institutions for kindly providing material and other assistance: Randall T. Schuh (AMNH), Mick Webb (BMNH), Dávid Rédei and András Orosz (HNHM), Eduardo Osuna (IZAV), Joseph C. Schaffner (TAMU), Steve Heydon (UCDC), and Thomas J. Henry (USNM). Special thanks are extended to Ernesto Barrera (UNAM) and Diana Martinez (UNAM) for the preparation of the pictures, and to Oscar Francke Ballve (UNAM) for his critical reading of the manuscript with helpful comments.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 405 H. BRAILOVSKY

References

BARANOWSKI R. M. & SLATER J. A. 2005: The Lygaeidae of the West Indies. University of Florida, IFAS. Florida Agricultural Experiment Station Bulletin 402: 1–266. BARBER H. G. 1939: Insects of Porto Rico and the Virgin Islands. Hemiptera-Heteroptera (excepting the Miridae and ). Scientific Survey of Porto Rico 14(3): 263–441. BARBER H. G. 1947: The family Lygaeidae (Hemiptera-Heteroptera) of the island of Cuba & the Isle of Pines. Memorias de la Sociedad Cubana de Historia Natural 19: 55–75. BARBER H. G. & ASHLOCK P. D. 1960: The Lygaeidae of the Van Voast-American Museum of Natural History expedition to the Bahama Islands, 1953. Proceedings of the Entomological Society of Washington 62: 117–124. DISTANT W. L. 1880–1893: Insecta. Rhynchota, Hemiptera-Heteroptera. In: GODMAN F. D. & SALVIN O. (eds.): Biologia Centali-Americana. Vol. 1. London, xx + 462 pp., 39 pls. FROESCHNER R. C. 1999: True Bugs (Heteroptera) of Panama: A synoptic catalog as a contribution to the study of Panamanian biodiversity. Memoires of the American Entomological Institute 61: 1–393. HENRY T. J. 1997: Phylogenetic Analysis of Family Groups within the infraorder Pentatomomorpha (Hemiptera: Heteroptera) with emphasis on the Lygaeoidea. Annals of the Entomological Society of America 90: 275–301. MONTANDON A. L. 1907: Hémiptères Hétéroptères. Espéces nouvelles ou peu connues. Buletinul Societãþii de ªtiinþe din Bucureºti, Româna 16: 292–295. SLATER J. A. 1964: A catalogue of the Lygaeidae of the World. Volumes 1 and 2. University of Connecticut, Storrs, 1688 pp.

406 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 407–413, 2013

A new species of Kanadyana (Hemiptera: Heteroptera: ) from Ghana

ELÕD KONDOROSY Department of Animal Science, Georgikon Faculty, University of Pannonia, Deák F. u. 16, Keszthely, H-8360 Hungary; e-mail: [email protected]

KONDOROSY E. 2013: A new species of Kanadyana (Hemiptera: Heteroptera: Rhyparochromidae) from Ghana. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 407–413. – A new species of the genus Kanadyana Scudder, 1963, Kanadyana stehliki sp.nov. (Hemiptera: Heteroptera: Rhyparochromidae: Rhyparochrominae: Cleradini) is described from Ghana. A male of Kanadyana dolora Scudder, 1963 is described and the species is recorded as new for the fauna of Ghana and Togo. A key to the species of the genus is provided. Keywords. Heteroptera, Rhyparochromidae, Kanadyana, true bugs, seed bugs, taxonomy, new species, new record, Afrotropical Region, Ghana, Togo

Introduction Cleradini is one of the smaller tribes of the family Rhyparochromidae, the seed bugs (Hemiptera: Heteroptera) and, with the exception the Neotropical Bathycles amarali (Correa, 1956), originally also assigned to this tribe but now included in tribe Udeocorini (SCUDDER 1962), the only known blood-sucking group of the infraorder Pentatomomorpha to feed on vertebrate hosts. This tribe has been reviewed by MALIPATIL (1983). The group’s original distribution covers the tropical regions of the eastern hemisphere and contains 19 genera and 54 valid species. Half of the known species live in Australia (as revised by MALIPATIL 1981), and the most recently discovered species have been recorded from there (MALIPATIL & BLACKET 2011) or from New Guinea (BRAILOVSKY 2011). The neighbouring Oriental Region is slightly richer in species than the Afrotropical Region. The only species occurring in America is Clerada apicicornis Signoret, 1863, the earliest described species of the group, and now introduced worldwide. The twelve known African Cleradini species belong to seven genera, all of which are endemic to the continent, except Navarrus Distant, 1901 with the single species, N. phaeophilus (Walker,1872) which inhabits also India and the Philippines (HARRINGTON 1983). The majority of Afrotropical taxa were discovered by SCUDDER (e.g. 1963, 1969). The genus Kanadyana Scudder, 1963 was described as monotypic, containing only K. dolora with the description based on three females from Congo and Cameroon (SCUDDER 1963). Later MALIPATIL (1983) added certain features and figured the female genitalia after study of the holotype; the male remained unknown. In this contribution, a further new species of Kanadyana is described from Ghana, the male of K. dolora is described, and new distributional data for the species are added.

407 E. KONDOROSY

Material and methods External and genital structures were studied with an Olympus SZ 11 stereoscopic microscope. The label data are cited verbatim with data on different rows separated by a slash (/) and data on different labels by a double slash (//); [hw] = preceding text handwritten, [p] = preceding text printed (only if part of the label handwritten).

The institutions in which the specimens are deposited are abbreviated as: HNHM ...... Hungarian Natural History Museum, Budapest, Hungary NHMW ...... Natural History Museum, Vienna, Austria NMPC ...... National Museum, Prague, Czech Republic

Kanadyana dolora Scudder, 1963 (Figs 1, 3, 4a, b, c)

Type material examined. Paratype: ♀, ‘Manengole / Cameroon / IV. 49 / R. Tesárek leg.’ (NMPC). Additional material examined. CAMEROON: 1 ♂ 1 ♀, ‘Manengole / Cameroon / IV. 49 / R. Tesárek leg.’ (NMPC). GHANA: 1 ♀, ‘Ghana – Central Reg. / Kakum forest N05° / 20′50″ W01°22′59″ // 135m, on light, 06–08. III. 2009 / leg. Kondorosy’; 1 ♂ 2 ♀♀, ‘Ghana – Central Reg. / Abrafo, Kakum forest, 2–13. X. 2008, leg. G. Csontos’; 1 ♂, ‘Ghana – Central Reg. / Kakum forest N05° / 21′17″ W01°22′19″ // 180m, on light, 06–18. IV. 2009 / leg. G. Csontos’. TOGO: 2 ♂♂, ‘Togo, Mt. Klouto, / Kouma-Konda / 22–24. IV. 2009 / leg. G. Csontos’ (HNHM, Kondorosy collection). Description of male. Coloration and basic body structure as in female (Fig. 1). In order to facilitate comparison with the newly-described species, the coloration is given here. Coloration. Head fuscous, antenna multicolorous: segment I dark brown, apically reddish, II yellow, III reddish-brown to fuscous, IV narrowly dark basally, its basal half mostly yellow changing to reddish in the apical half. Pronotum dark brown except for pale posterior margin. Scutellum fuscous. Clavus brown, emergent vein AA1+2, narrow apical and lateral margin ochraceous; corium brown, emergent vein Cu, most parts of basal half, small spots on apical half and exocorium ochraceous; membrane pale brown, with translucent veins and apex, and with dark brown at mid-base and in inner angle. Dark coloration of corium slightly variable but almost always continuous along costal margin, leaving an isolated tiny spot on vein R. Legs and labium yellow. Ventrally brown, medial part paler. Male genitalia. Pygophore opening with straight dorsal margin (Fig. 3), laterally with obtuse edge. Paramere relatively short, apical arm widened at basal part, afterwards narrowing to acute apex (Fig. 4b), whole strongly incurving for entire length; inner arm short, pointing inwards at almost right-angles (Fig. 4c); shaft for most of its length with a short, broad, lamellate projection ending in two blunt tips (Fig. 4a); no third arm in inner surface. Measurements (n = 5 ♂♂, 5 ♀♀; the first values are for the males, the second for females, all measurements in mm; for labial segments only 1 ♂ and 1 ♀ measured). Body length: 7.2–7.6, 8.8–9.3. Head: length 1.27–1.44, 1.56–1.65; maximum width across eyes 1.05–1.11, 1.14–1.28; interocular space 0.35–0.39, 0.40–0.46; eye length 0.50–0.52, 0.55–0.61. Lengths of antennal segments: I – 0.53–0.59, 0.65–0.71; II – 1.12–1.23,

408 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Kanadyana

Figs 1–2. Kanadyana spp., habitus. 1 – K. dolora Scudder, 1963; 2 – K. stehliki sp.nov.

1.24–1.33; III – 0.61, 0.68–0.71; IV – 1.31, 1.48–1.64. Length of labium: 3.10, 3.53; labial segments I – 0.70, 0.78, II – 0.57, 0.68, III – 1.32, 1.50, IV – 0.51, 0.57. Pronotum: length 1.12–1.25, 1.38–1.54; maximum width 2.05–2.19, 2.54–2.61. Scutellum length 0.97–1.11, 1.25–1.40. Claval commissure length 0.94–1.01, 1.26–1.35. Distribution. Cameroon, Congo (SCUDDER 1963), Ghana, Togo (new records).

Kanadyana stehliki sp.nov. (Figs 2, 5, 6a, b, c)

Type material. Holotype: ♂ (HNHM), ‘GHANA, Eastern Region / Atewa Range, 730 m / 06°13′50″N 0°33′27″W // at light / 26–28.III.2009 / leg. E. Kondorosy’. Paratypes: 1 ♀, same data as holotype (HNHM); 2 ♂♂ 1 ♀, ‘GHANA-Eastern Reg. / Atewa Range, N06° / 13′56″ W0°33′07″ / 660 m, on light / 19–23.III.2009 / leg. Kondorosy’ (1 ♂ NMPC, 1 ♂ 1 ♀ Kondorosy collection); 2 ♀♀, ‘GHANA-Eastern Reg. / Atewa Range, N06° / 13′50″ W0°33′27″ / 730 m, on light / 26–28.III.2009 / leg. Kondorosy’ (HNHM, NHMW). Description. Coloration. Body yellowish-brown (Fig. 2). Head brown to fuscous, apex of antennal segment III and very base of IV also brown. Anterior half of pronotum concolorous with head. Basal half of scutellum also brown occasionally. Clavus dark brown between inner and middle rows of punctures and on outer row. Corium dark brown on both inner rows of punctures, on lateral row and in apical half except exocorium and

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 409 E. KONDOROSY a subapical spot connected with it. Pale parts of clavus and endocorium yellowish-white. Membrane transparent, with a narrow triangular spot at inner angle basally and dusky stripes between veins apically. Most parts of thoracic sterna dark brown. Punctures very dense and minute on head, interrupted by two large, very finely- wrinkled spots between eyes. Anterior half of pronotum punctate in similar fashion to head, leaving two finely-wrinkled elevations almost free of punctures; posterior half with punctures much larger and on disk relatively sparse (laterally very dense), posterior margin and slightly elevated median carina smooth. Punctures of scutellum and hemelytra similar to, or slightly finer than, those on posterior pronotal half, on scutellum leaving rear half beyond median impunctate. Clavus with three complete rows of punctures, posterior half with many further punctures between two inner rows. Endocorium with two regular rows of punctures along clavus, corium between Sc and M densely punctate, posterior half also with punctures between M and Cu. Punctures of thoracic sterna finer than those of posterior pronotal half, abdomen almost impunctate, strongly transversely wrinkled. Structure. Body lustrous, with short, quite dense, upright hairs (slightly shorter than diameter of tibiae); some longer hairs among them. Legs, especially fore-femora, with very long hair-like setae, tibial setae not thicker than normal hairs. Legs unarmed, fore- femora not thicker than others. Male genitalia. Pygophore opening with arched dorsal margin, laterally rectangular (Fig. 5). Paramere large, in resting position nearly reaching opposite margin of pygophore opening; apical part long, parallel (Fig. 6b), bending inwards slightly, with blunt apex; inner arm broad, strong, curved in apical half (Fig. 6c); shaft without lamellate projection but with a long, acute, spine-like projection near base of arms (Fig. 6a). Measurements (the first values are for males, the second for females, all measurements in mm; for labial segments only 1 ♂ and 1 ♀ measured). Body length: 7.1–7.2, 8.2–9.0. Head: length 1.37–1.41, 1.44–1.60; maximum width across eyes 1.05–1.13, 1.18–1.25; interocular space 0.32–0.35, 0.43–0.50; eye length 0.49–0.51, 0.52–0.55. Lengths of antennal segments: I – 0.55–0.60, 0.60–0.65; II – 1.02–1.06, 1.10–1.18; III – 0.48–0.53, 0.55–0.59; IV – missing, 1.58–1.60. Labium length 3.22, 3.42, labial segments I – 0.68, 0.72, II – 0.61, 0.63, III – 1.42, 1.49, IV – 0.51, 0.57. Pronotum: length 1.12–1.18, 1.37–1.52; maximum width 1.98–2.04, 2.44–2.63. Scutellum length 1.00–1.06, 1.18–1.30. Claval commissure length 0.95–1.02, 1.17–1.31. Variability is slight. Sometimes antennal segment I is slightly darker than II and/or segment IV is brownish in apical part. Posterior half of pronotum sometimes only slightly paler than anterior part, and pronotal median carina sometimes not at all elevated. Differential diagnosis. Kanadyana stehliki sp.nov. differs from K. dolora, the only other known Kanadyana species, most strikingly in the very different paramere (cf. Figs 4 and 6). Difference in coloration is also important, but there are transitions in some features, as in pronotal and corial coloration. Therefore the only reliable external difference is the colour of the antenna: in K. stehliki sp.nov. all the segments are pale, while K. dolora has antennal segments I and III mostly dark (apical parts can be paler).

410 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 New species of Kanadyana

Figs 3–6. Kanadyana spp., pygophore, dorsal view (3, 5) and right paremere (4, 6): a – ventral view, b – dorsal view, c – caudal view. 3, 4 – K. dolora Scudder, 1963; 5, 6 – K. stehliki sp.nov.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 411 E. KONDOROSY

Etymology. I dedicate this species to Dr. Jaroslav L. Stehlík for the occasion of his 90th birthday and in honour of his enormous contributions to heteropterology, especially his knowledge of Pyrrhocoroidea. Distribution and habitat. All the specimens were collected in Ghana in the Atewa Mountains, and distribution appears limited to this isolated area. The surrounding areas were once rain forests but these have largely fallen victim to human settlement, agriculture, mining and logging and only small fragments survive. Otherwise, these mountains are unique, with upland evergreen forests and other rare biotopes, and many rare and endemic species (such as the pierid butterfly Mylothris atewa Berger, 1980) (RAP 2007). All the known Kanadyana stehliki sp.nov. specimens were collected at light, relatively high above sea level. The biology is otherwise unknown.

Key to the species of Kanadyana 1 Antenna multicolorous, segment I dark brown, apically reddish, II yellow, III reddish-brown to fuscous, IV narrowly dark basally, basal half mostly yellow changing to reddish in apical half; dark coloration of corium reaching costal margin (Fig 1); paramere short, apically narrowing to pointed tip, lateral arm short, strongly hooked inwards (Figs 4a–c)...... K. dolora Scudder, 1963 – Antenna almost unicolorous, yellow; dark corial spot not reaching costal margin (Fig 2); paramere long, apical part parallel, lateral arm long, hooked only apically (Figs 6a–c)...... K. stehliki sp.nov.

Acknowledgements I am indebted to Petr Kment (NMPC) for the opportunity to study the paratype of K. dolora and to him and Dávid Rédei (HNHM) for the loan of the Kanadyana specimens from their museums. I am very grateful to Claudia Molnár for revising the text, and to Katinka Varga for taking the pictures (both University of Pannonia).

References

BRAILOVSKY H. 2011: A new species of Pactye from Papua New Guinea (Hemiptera: Heteroptera: Rhyparochromidae). Acta Entomologica Musei Nationalis Pragae 51: 443–448. HARRINGTON J. B. 1983: A new species of Cleradini (Hemiptera: Lygaeidae: Rhyparochrominae) from the Central African Republic and Ghana. Journal of the New York Entomological Society 91: 63–67. MALIPATIL M. B. 1981: Revision of Australian Cleradini (Heteroptera: Lygaeidae). Australian Journal of Zoology 29: 773–819. MALIPATIL M. B. 1983: Revision of World Cleradini (Heteroptera: Lygaeidae), with a cladistic analysis of relationships within the tribe. Australian Journal of Zoology 31: 205–225. MALIPATIL M. B. & BLACKET M. J. 2011: Three new species of Cleradini from Australia (Hemiptera: Heteroptera: Rhyparochromidae). Zootaxa 3003: 43–54. RAP (RAPID ASSESSMENT PROGRAM) 2007: Biodiversity in the Atewa Range Forest Reserve, Ghana. Conservation International, Arlington, VA, USA. 28 pp.

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SCUDDER G. G. E. 1962: The World Rhyparochrominae (Hemiptera: Lygaeidae) I. New synonymy and generic changes. Canadian Entomologist 94: 764–773. SCUDDER G. G. E. 1963: The World Rhyparochrominae (Hemiptera: Lygaeidae) III. New Rhyparochrominae from the Ethiopian Region. Canadian Entomologist 95: 1233–1253. SCUDDER G. G. E. 1969: The World Rhyparochrominae (Hemiptera: Lygaeidae) IX. Four new genera collected by Dr. R. Linnavuori in the Sudan. Annales Entomologici Fennici 35: 85–94.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 415–432, 2013

Carduelicoris stehliki, a new genus and species of (Hemiptera: Heteroptera) from Madagascar

PETR KMENT Department of Entomology, National Museum, Kunratice 1, CZ-148 00 Praha 4, Czech Republic; e-mail: [email protected]

KMENT P. 2013: Carduelicoris stehliki, a new genus and species of Pentatomidae (Hemiptera: Heteroptera) from Madagascar. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 415–432. – Carduelicoris stehliki gen. et sp.nov. (Hemiptera: Heteroptera: Pentatomidae: Pentatominae) is described from Madagascar and placed in the tribe Halyini. The new genus differs markedly from all the remaining Madagascar representatives of the tribe by the produced, apically dentate humeral angles of the pronotum. The new taxon is dedicated to Jaroslav L. Stehlík, the doyen of the Czech and Slovak heteropterology, on the occassion of his 90th birthday. Diagnostic characters of the tribe Halyini are discussed. Keywords. Heteroptera, , Halyini, new genus, new species, taxonomy, Madagascar

Introduction The fauna of the superfamily Pentatomoidea (Hemiptera: Heteroptera) of Madagascar was monographed by CACHAN (1952), covering all families except . Madagascan Pentatomoidea received very little attention during the next fifty years, most of which was only a minor component of more widely conceived research. Concerning taxonomy, nomenclature and distribution of Pentatominae, LESTON (1953a), SCHOUTEDEN (1954), DAY (1965), ORIAN (1965), GREATHEAD (1968), LINNAVUORI (1973, 1975), COUILLOUD (1989), AHMAD (1995), RIDER (1998a, b, 2007), GÖLLNER-SCHEIDING (1999) and ŠTYS & EXNEROVÁ (2003) provided contributions on various topics. A few revisions devoted solely or partly to Madagascan taxa were published in recent years, documenting the reborn interest in the highly endemic heteropteran fauna of the island (KMENT 2008, 2011, 2012; KMENT & JINDRA 2009; FERRARI et al. 2010). Finally, ARNOLD (2011) attempted to erect a new genus, Jostenicoris, to accommodate the Afrotropical and Madagascan species of the Palaeotropical genus Agaeus Dallas, 1851 and placed in the tribe Strachiini, while the genus Agaeus s. str. as recognized by him is limited to the Oriental Region and in his opinion it belongs to the tribe Halyini. However, as ARNOLD (2011) failed to provide any description or definition of his new genus, Jostenicoris Arnold, 2011 remains unavailable name (see ICZN 1999: 13.1.1) and must be listed as a junior synonym of Agaeus being a nomen nudum. In this paper I provide description of an additional new genus and species, one of the many necessary steps for compiling an updated monograph of Pentatomidae of Madagascar.

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Material and methods In quoting the labels of the type specimens, a slash (/) is used to divide data on different rows of one label, a double slash (//) is used to divide data on different labels, authors’ comments are given in square brackets [], and the following abbreviations are used: hw = handwritten, p = printed. Unless stated otherwise, the described labels are white. The specimens examined are deposited in the following collections: BMNH ...... Natural History Museum, London, United Kingdom MNHN ...... Muséum national d’Histoire naturelle, Paris, France MMBC ...... Moravian Museum, Brno, Czech Republic NMPC ...... National Museum, Praha, Czech Republic

Measurements were taken from dry-mounted specimens under a MBS-10 stereomicroscope using an ocular micrometer. The following dimensions were measured: body length (from apex of mandibular plates to apex of membrane), head length (from apex of mandibular plates to anterior margin of pronotum), head width (maximum width across eyes), interocular width (between inner margins of compound eyes), length of each antennal segment (maximum length), pronotum length (medially in most exposed, anterodorsal view), pronotum width (maximum width between processes on humeral angles), scutellum length (medially from base to apex), scutellum width (maximum width at base), and abdomen width (maximum width across hypopleurites III). Dissections of genitalia and all line drawings were made under a Leica MZ75 stereomicroscope with a camera lucida. Non-coated specimens were examined by Hitachi S-3700N environmental electron microscope at the Department of Paleontology, National Museum in Prague. Habitus photographs were taken using a Canon MP-E 65 mm macro lens attached to a Canon EOS 550D camera and stacked from multiple layers using the Helicon Focus 5.1 Pro software. For the study of genitalia, specimens were softened in distilled water, and the male pygophore or female abdomen were removed under the stereomicroscope using sharp forceps, then put into concentrated solution of KOH and heated (but not as much as to boil). After the KOH treatment, the pygophore or abdomen were washed in distilled water and dissected under the stereomicroscope. The dissected phallus and female genitalia were subsequently stored in plastic microvials with glycerol attached to the same pin as the specimen. The general morphological terminology follows mostly TSAI et al. (2011); parts of the thoracic scent efferent system of the metathoracic scent glands are named in accordance with KMENT & VILÍMOVÁ (2010) and those of pygophore according to SCHAEFER (1977).

Results Carduelicoris gen.nov. Type species. Carduelicoris stehliki sp.nov., by present designation. Description. Body mostly parallel-sided with strongly prominent humeral angles (Figs 1–3).

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Figs 1–6. Carduelicoris stehliki gen. et sp.nov. 1–3 – habitus of the male holotype (body length 15.4 mm) in dorsal (1), ventral (2) and lateral (3) view; 4 – head in ventral view (magnification 23×); 5 – meso- and metapleuron with the external scent efferent system of the metathoracic scent gland (magnification 30×); 6 – detail of peritreme and mycoid microsculpture of evaporatorium (magnification 250×). Scale bars: 1 mm (4, 5), 0.1 mm (6).

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Head parallel-sided, compound eyes relatively small, conically protruding from head outline (Figs 1–2, 4, 7). Clypeus much shorter than mandibular plates, apically free (Fig. 7). Mandibular plates strongly, subtriangularly produced anteriad of clypeus, obtusely rounded apically, anterolaterally with one additional subapical, pointed tooth- like projection, divided from the apical portion by a narrow U- to V-shaped incision (Figs 1, 4, 7). Ocelli small, situated much posteriad of eyes, at the anterior margin of pronotum (Fig. 7). Antenniferous tubercle small, greatly concealed by mandibular plates, therefore only lateral extremity visible in dorsal view (Figs 1, 7). Antennae slender, antennal segments straight, narrowly cylindric, IIa, IIb and III slightly wider apically (Fig. 1); antennal segment I shortest and stoutest; lengths of antennal segments: I < IIa ≤ IV < IIb = III. Bucculae long and narrow, anteriorly truncate, anteroventrally sharply rectangular, posteriorly narrowed and slightly rounded (Fig. 8). Rostrum 4-segmented, reaching between metacoxae; lengths of rostral segments IV < I < II = III; segment I concealed by bucculae in rest (Fig. 4). Pronotum transversely subhexagonal, with strongly produced humeri (Figs 1–2, 9). Anterior margin as wide as head across eyes, concave, embracing postocular portion of head. Anterolateral angles prominent, directed anteriad, apically truncate, approaching posterior surface of compound eyes, anteriorly with denticle directed laterad. Anterolateral margins of pronotum deeply, roundly concave, not carinate, bearing 6–11 small callose denticles of variable shape (also varying between left and right sides of pronotum, see Fig. 9) distributed from anterolateral angle to the first triangular lobe of humeral angles. Humeri produced into large, lobe-like projection, projecting distinctly anteriad of eye, apically widely rounded and flattened, with four projections, the first (anterior) one narrowly triangular, acute, the second (apical) one narrowly lobate and apically rounded, the third and fourth (posterior) ones short, broadly lobate and rounded; pronotal margin between second and third lobe shallowly concave to nearly straight (Fig. 9). Lateral margins of pronotum weakly converging posteriad of humeral projections towards base of corium, there forming an obtuse angle and continuing obliquely towards base of scutellum; posterolateral angle obtuse; posterior margin transversal, straight. Pronotal disc highest along its posterior margin and towards apices of humeral angles, rather steeply sloping towards anterior margin (Fig. 3); pronotal disc anteriorly with two longitudinal depressions leading from cicatrices approximately towards anterolateral angles, diverging anteriad (Figs 1, 9). Scutellum (Fig. 1) triangular, lateral margins in frenal (= anterior) portion nearly straight, regularly narrowing towards apices of clavi, there slightly concave; postfrenal (= apical) portion of scutellum long, almost parallel-sided, broadly rounded apically. Scutellar disc elevated basally (Fig. 3), sloping laterad and posteriad, apical portion flat. Hemelytra (Fig. 1) reaching or slightly surpassing apex of abdomen. Clavus narrow. Corium laterally slightly widening towards segment III, then slightly narrowing towards apex, reaching middle of hypopleurite (= outer laterotergite) VI in rest; apical angle of corium narrowly rounded. Membrane widely rounded apically, veins subparallel. Thoracic pleura and sterna. Mesosternum anteriorly with short, wedge-shaped keel narrowing posteriad; mesosternum posteriorly and metasternum flat, not grooved. Ostiole of metathoracic scent glands situated in inner third of metapleuron width (Fig. 5), drop-

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Figs 7–10. Carduelicoris stehliki gen. et sp.nov. 7–8 – head: 7 – dorsal view (male from Sandringato); 8 – lateral view (female from Périnet). 9 – pronotum (male holotype, antero-dorsal view). 10 – paramere. Scale bars: 1 mm (7, 8, 9), 0.5 mm (10).

shaped, facing laterad, accompanied with short spout (about 2–3 times of ostiole length), narrowly rounded apically, directed laterad, with peritremal surface facing posteriad, with narrow median furrow in its basal half (Fig. 6); vestibule gibbose, elevated above surrounding pleuron; vestibular scar missing; periostiolar depression not developed; metapleural evaporatorium large, covering anterior and central parts of metapleuron, narrowing and reaching mesad between meso- and metacetabulum and laterad along metathoracic spiracle; mesopleural evaporatorium small, limited to postero-mesal portion of mesopleuron between metathoracic spiracle and mesopleural sulcus (Fig. 5); gyrification of metapleural evaporatorium well-developed, mesopleural evaporatorium mostly flat; metepimeral pseudosuture developed; metathoracic spiracle long and narrow (Fig. 5).

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Legs. Femora narrowly clavate, widest subapically, unarmed. Tibiae narrowest medially, slightly widening towards base and apex; lateral surface of tibiae sulcate. Tarsi 3-segmented, tarsomere 2 shortest. Abdomen widest across posterolateral angles of laterotergite III (Figs 1–2). Connexivum mostly not covered by corium, posterolateral angles of laterotergites obtusely rectangular, slightly protruding from the body outline (Fig. 1). Sternite III medially with two low longitudinal tubercles divided from each other by a shallow groove; remaining sternites evenly convex. Male genitalia. Pygophore (Figs 11–13) approximately as high (2.0 mm) as wide (2.0 mm); ventral wall slightly constricted under ventral rim in lateral view (Fig. 12). Ventral rim widely and shallowly concave, medially with deep and narrow U-shaped incision (Figs 11, 13). Ventral rim infolding wide and conspicuous, medially concave, dorsally continuing in a large, conical, dorsolaterally directed projection of ventral plus lateral rims (Figs 11, 13, 15). Lateral rim ridge-like, both lateral rim and lateral rim infolding fused with ventral rim and its infolding. Dorsal rim simple, medially not infolded, forming a rhomboid extension (Fig. 11); dorsal rim infolding reduced, producing articulated dorsal processes transversely elongate, mesally subtruncate, laterally rounded and bent caudally (Figs 11, 13, 15); outer surface of dorsal plate with specific mesh-like microsculpture (Fig. 16). Paramere sockets not developed. Parameres (Figs 10, 14) with basal apodeme large and eliptic, apical part of paramere flattened, hook-shaped, directed dorsad within pygophore at rest (Fig. 11), apex truncated, bearing scale-like microsculpture laterally (Figs 11, 14). Phallus (Figs 17–22) with phallotheca short and thick, barrel-shaped, strongly sclerotized, with distinctly protruding hinge; conjunctiva with two pairs of dorsal processes: first pair (dcp1) elongate, laterally sclerotized, second pair (dcp2) membranous, fused along midline into a single short, bulb-shaped lobe provided with two small dorsal projections; a pair of ventral conjunctival processes (vcp) (= median penial plates sensu GROSS 1975) sclerotized, dorsally fused, ventrally produced into a pair of large and broad lobes, embracing a thin tubular vesica curved ventrad, hardly visible between lobes of vcp; endophallic reservoir restricted approximately to apical two-thirds of inner lumen of phallotheca. Female genitalia. External structures. Terminalia as in Figs 23–24. Valvifers VIII (vf8) medially straight, adjacent to each other, laterotergites IX (lt9) directed obliquely at rest, basally parallel-sided, apically nearly parabolic; valvifers IX (vf9) well-exposed at rest, trapezoid, with a pair of elongate, basolaterally directed arms; triangulin missing. Genital tracts (Fig. 25) partly desintegrated during dissection. Gynatrium with a pair of small, well-distinguishable ring sclerites (Figs 24–25: rs) and a pair of additional, elongate sclerites situated caudad of ring sclerites; spermathecal opening (Figs 24–25: so) associated with an elongate sclerite (arcus, Figs 24–25: arc). Spermatheca (Fig. 25) with proximal duct relatively long, somewhat shorter than distal invagination of spermathecal duct, thin; spermathecal dilation (sd) large, spherical, inside milky, not transparent (possibly artefacts); distal invagination of spermathecal duct (‘sclerotized rod’) robust, basally and apically widened; intermediate part (ip) straight and narrow, provided with proximal (pfl) and distal flanges (dfl), apical receptacle (ar) small, spherical, bearing two processes (Fig. 25).

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Figs 11–16. Carduelicoris stehliki gen. et sp.nov. 11–13 – pygophore: 11 – intact, dorso-posterior view (magnification 45×); 12 – lateral view (magnification 47×); 13 – dissected, posterior view (magnification 50×). 14 – paramere (magnification 120×). 15–16 – dorsal plate of pygophore in situ (15, magnification 75×) and detail of microsculpture (16, magnification 210×). Scale bars: 0.5 mm (11, 12, 13, 15), 0.2 mm (14, 16).

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Differential diagnosis. The new genus differs from all other Madagascan Pentatomidae by combination of the shape of head (anteocular spine missing; clypeus distinctly shorter than mandibular plates but apically free; mandibular plates parallel-sided laterally, produced and narrowly rounded apically, with one subapical lateral tooth) and pronotum with protruding humeral angles curved anterolaterad, apically widely rounded and bearing small lobes and denticles on their margin. The male genitalia are peculiar within Halyini by presence of dorsal plates of pygophore. See also Discussion below. Etymology. The generic name is composed of the Latin noun carduelis, -is, meaning “Goldfinch” in English and “stehlík” in Czech (viz also Carduelis carduelis Linnaeus, 1758) and the Latinized Greek noun coris, meaning true bug; gender is masculine. The name is dedicated to Jaroslav L. Stehlík, the doyen of the Czech and Slovak heteropterology, on the occassion of his 90th birthday. Included species. Monotypic.

Carduelicoris stehliki sp.nov. (Figs 1–26) Type locality. East Madagascar, Sandrangato (at road from Moramanga to Anosibe an’Ala), 19°06′15″S 48°14′36″E (see Fig. 26). Type material. Holotype: ♂ (Figs 1–3), ‘Sandrangato [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // Brit.Mus. / 1965-338 [p] // ♂ [p] // Genus not / in B. M. [hw] / W.E.China det. 196[p]4[hw] // HOLOTYPUS / CARDUELICORIS / STEHLIKI / sp. nov. / det. P. KMENT 2013 [p, red label]’ (BMNH) (the specimen pinned through pronotum, left hind leg and right antennal segments III and IV missing). Paratypes (3 ♂♂ 2 ♀♀): ♂, ‘Sandrangato [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // MUSEUM PARIS [p] // ♂ [p]’ (MNHN); ♂, ‘Sandrangato [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // MUSEUM PARIS [p] // ♂ [p] // Genus not / in B. M. [hw] / W.E.China det. 196[p]4[hw]’ (MNHN) (detached pygophore glued on a piece of card attached to the same pin); ♂, ‘Sandrangato [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // ♂ [p]’ (NMPC) (dissected pygophore and parameres glued on a piece of card and phallus stored in a small plastic microvial with glycerol attached to the same pin); ♀, ‘Sandrangato [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // Brit.Mus. / 1965-338 [p] // ♀ [p]’ (BMNH); ♀, ‘Sandrangato [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // MUSEUM PARIS [p] // ♀ [p]’ (MNHN) (abdominal dorsum, venter and antennomere IV glued on a piece of card, dissected female genitalia placed in a plastic microvial with glycerol attached to the same pin); ♀, ‘Périnet [p] // INSTITUT / SCIENTIFIQUE / MADAGASCAR [p] // MUSEUM PARIS [p] // ♀ [p]’ (MNHN). Each paratype bearing the following identification label: ‘PARATYPUS / CARDUELICORIS / STEHLIKI / sp. nov. / det. P. KMENT 2013 [p, red label]’; 1 ♂ 1 ♀, ‘MADAGASCAR, 2007, / Lakato env., near Andasibe-Mantadia N.P., Z. Mráèek leg., 9.–10.i. [p] // ‘♂’ or ‘♀’ [p] (MMBC). Description. Coloration beige, head posteriorly and pronotum anteriorly tinged with yellow; femora, base of abdomen, and apices of humeral angles brown (Figs 1–2); apical portions of tarsal claws and abdominal spiracles black; abdominal dorsum and ventral side of hemelytra beige; membrane translucent, very slightly infumate, with brown veins. Vestiture. Body glabrous; antennal segments IIb and III apically with short and sparse setae, entire antennal segment IV densely covered with short and fine pale setae intermixed with somewhat longer and stouter brownish setae; labium ventrally, ventral surface of femora and basal portion of tibiae with sparse short pale setae; tibiae apically and tarsi densely covered with longer and stouter brownish setae (Fig. 4). Body dorsally covered with small, shallow punctures, mostly concolorous beige, but sometimes brown

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Figs 17–22. Phallus of Carduelicoris stehliki gen. et sp.nov. 17–19 – intact, 20–22 – inflated; 17, 20 – dorsal view, 18, 21 – lateral view, 19, 22 – ventral view. Abbreviations: dcp1–2 – dorsal conjuctival processes 1 and 2, er – endophallic reservoir, vcp – ventral conjuctival processes, ve – vesica. Scale bar: 0.5 mm.

or black. Two longitudinal depressions anteriorly on pronotum with bottom densely covered with black punctures (Figs 1, 9), adjacent surface between and laterad of the depressions slightly raised, callous and sparsely punctured. Scutellar disc laterally on each side with two more or less apparent longitudinal, callous, raised lines, posteriorly merging with a single, wide median longitudinal callosity occupying about one median third of scutellum length (Fig. 1); apex of scutellum not callous. Clavus basally with 5 rows of punctures, anterior four of them rather irregular, continually merging into a single row reaching claval apex; the posteriormost (fifth) row along posterior claval margin ending approximately in claval mid-length. Corium along claval fold and cubital vein (Cu) with slightly raised unpunctured longitudinal lines, surrounding punctures organised in more or less regular lines; corium medially (between claval fold and Cu) with irregularly distributed punctures and unpunctured areas (Fig. 1). Posterolateral angles of laterotergites with minute yellowish callosities. Body ventrally irregularly punctured, punctures shallow to deep, concolorous to black; bases of mesopleura, metapleura and sternites III and IV medially with deep black punctures (Fig. 2). No waxy secretion was observed anywhere on the body.

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Figs 23–25. Female genitalia of Carduelicoris stehliki gen. et sp.nov. 23–24 – external female terminalia (23 – intact, 24 – dissected); 25 – internal female genitalia. Abbreviations: ar – apical receptacle of spermatheca, arc – arcus, dfl – distal flange, ip – intermediate part of spermatheca, lt8–9 – laterotergites VIII–IX, pd – proximal duct of spermatheca, pfl – proximal flange, rs – ring sclerite, sd – spermathecal dilation, so – sclerite surrounding spermathecal orifice, sr – sclerotised rod, t8 – tergite VIII, vf8–9 – valvifers VIII–IX, x – segment X. Scale bars: 1 mm.

Measurements (in mm; median (minimum–maximum)). Males (n = 4, for antennal segments: II – n = 3, III and IV – n = 2): Body length 16.4 (15.7–16.5); head length 3.53 (3.37–3.58); head width 3.37 (3.37–3.47); interocular width 2.25 (2.15–2.25); lengths of antennal segments: I – 0.97 (0.92–1.02), IIa – 1.33 (1.33–1.43), IIb – 1.94 (1.94–2.0), III

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– 1.94–2.04, IV – 1.43–1.53; pronotum length 3.58 (3.47–3.68); pronotum width 9.45 (9.20–9.50); scutellum length 5.57 (5.31–5.62), scutellum width 4.34 (4.29–4.60); abdomen width 8.43 (8.18–8.58). Females (n = 3, for antennal segment III n = 1, antennal segment IV missing in all specimens): Body length 17.1 (16.6–17.8); head length 3.68 (3.68–3.78); head width 3.47 (3.47–3.47); interocular width 2.35 (2.25–2.35); lengths of antennal segments: I – 0.92 (0.92–1.02), IIa – 1.33 (1.33–1.43), IIb – 1.94 (1.94–1.94), III – 1.94; pronotum length 3.68 (3.58–3.68), pronotum width 9.61 (9.50–10.32); scutellum length 5.72 (5.52–6.23); scutellum width 4.80 (4.50–4.91); abdomen width 8.89 (8.79–9.30). Variability. Males and females do not differ externally except for terminalia; females seem to be slightly larger than males. Regardless of the sex, the specimens differ in shape and number of denticles on the anterolateral margins and humeral angles of the pronotum, and in the shape of the apical lobes of the humeral angles (e.g., the margin between the second and third lobe is slightly concave to straight), which are usually also variable between the left and right sides of the body (Fig. 9). There are also slight differences in the pattern of callosities and punctures on both dorsal and ventral surfaces of the body. Etymology. The new species is dedicated to Jaroslav L. Stehlík. Biology. Unknown. Distribution. Known only from three localities in eastern Madagascar in the zone of tropical rain forests, Sandrangato (the type locality), ,Périnet (18°55′39″S, 48°24′52″E), and Lakato (19°11′S, 48°26′E (Fig 26).

Discussion Tribal placement of Carduelicoris gen.nov. Using the key to tribes of Madagascar Pentatominae by CACHAN (1952), Carduelicoris gen.nov. belongs to ‘groupe II’ of tribes including Nealeriini, Memmiini, Phricodini, Halyini, Amyntorini, Triplatygini and Rolstoniellini (= Compastini), being characterised mostly by shape of the head and absence of a tubercle at base of abdomen. Within ‘groupe II’, CACHAN (1952) placed the genera with 2-segmented tarsi in the tribe Nealeriini, and those with 3-segmented tarsi and 4-segmented antennae in the Memmiini and Phricodini. The remaining genera, sharing 3-segmented tarsi and 5-segmented antennae (the most common combination of characters in Pentatomidae), were divided among four tribes (Halyini, Amyntorini, Triplatygini and Rolstoniellini). Although Halyini is one of the best known taxa within Pentatominae, diagnosis of this group is very problematic (cf. GROSS 1976: 252). In most of treatments, Halyini are defined based on few morphological characters in reference to the local fauna under study (e.g. DISTANT 1902, CACHAN 1952, ROLSTON & MCDONALD 1979, ABBASI 1986, DERJANSCHI & PÉRICART 2006, MEMON et al. 2011), but none of them apply for all the known genera included in this taxon. GROSS (1976) addressed this problem in detail, giving the following external characteristics of the tribe: mostly large species with

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 425 P. K MENT elongate body-shape; characteristic shape of head (mandibular plates sometimes produced in front of the clypeus as spinose or irregularly truncate lobes or not; antenniferous tubercles in some species armed exteriorly with a hook-like process, in others not); antennae 5-segmented (in some genera 4- or even 3-segmented); pronotum hexagonal, with humeral angles frequently armed (sometimes as spines, sometimes as horn-like processes); scutellum always triangular (but large and reaching apex of abdomen in Afrotropical Mezessea Linnavuori, 1982: see LINNAVUORI 1982); hemelytra and hind wings fully developed (except Australian Tinganina Bergroth, 1909); hypopleurites frequently strongly spined posteriorly; rostrum sometimes very long, almost reaching the apex of the abdomen, only rarely (if ever) failing to reach base of abdomen; abdominal venter often with longitudinal groove to receive the long rostrum. In addition, DISTANT (1902) and ABBASI (1986) pointed out the head distinctly longer than broad (though sometimes its length being subequal to the width) and prominent eyes, and DISTANT (1902) and DERJANSCHI & PÉRICART (2006) noted that the antennal insertions are separated from the eyes by a distance at least equal to eye diameter. The shape of the peritreme of the metathoracic scent gland is also considerably variable within the tribe (see e.g. CACHAN 1952 and LINNAVUORI 1982). LESTON (1953b) suggested that the tribe Halyini might be defined on the basis of presence of wax glands. Most of the characters mentioned above are quite weakly defined and they are difficult to use in deciding whether a particular species belongs to Halyini or not. The shape of the head is usually elongate in the tribe, but the genera are too diverse in respect of this character to make a clear-cut definition. Several characters (5-segmented antennae, hexagonal pronotum, triangular scutellum, fully developed wings) are clearly plesiomorphic within Pentatomidae and occur in the majority of the other tribes too. Most of the apparent apomorphies (spined lateral margin of the abdomen, long labium, grooved abdominal venter) occur sporadically in several other tribes as well. Wax glands are also not unique to the halyines (ROLSTON & MCDONALD 1979). In spite of these problems, still Halyini apparently forms a clade of taxa of quite similar general habitus, but they are difficult to define based on external characters. According to GROSS (1976), Halyini can be well-defined only on the base of male genitalia, especially the phallus: the pygophore strongly tends to be laterally produced into a pair of robust or acute spinous or horn-like processes directed posteriad, sometimes subdivided (in other species the lateral lobes are only incipient or truncate); the phallus is always with strongly sclerotized phallotheca (= phallosoma sensu GROSS 1975) which is generally quite small in size in relation to conjunctiva, and in many taxa with transparent apical thecal shield surrounding the dorsal opening of the theca; conjunctiva large, frequently distinctly longer than the theca and very frequently with 2 or 3 pairs of conjunctival processes present, some of them can be secondarily subdivided or fused along midline in other taxa (see also MEMON et al. 2011); the ventral conjunctival processes (= median penial plates sensu GROSS 1975) small and directed downwards, only their ventral apices are usually external to the conjunctiva, sometimes adjacent or even fused ventrally; the gonopore always opens between the ventral conjunctival processes and the vesica does not protrude beyond them. On the other hand, parameres

426 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Carduelicoris stehliki gen. et sp.nov. of Pentatomidae vary a great deal within the tribe (GROSS 1976, MEMON et al. 2011) and can be even reduced (in Solomonius group, see WALL 2007). In GROSS’s (1976) opinion, the female spermatheca of Halyini is of typical pentatomid pattern, its apical receptacle bearing two short tubular processes. However, the apical receptacle lacks any processes in Carenoplistus Jakovlev, 1882 and Lodosocoris Ahmad et Afzal, 1986 (MEMON et al. 2011) while other genera have three, four or even a larger number (up to 16) of processes (Paranevisanus Distant, 1908: see MEMON et al. 2011). Moreover, MEMON et al. (2006) documented the infraspecific variability of the shape and number of spermathecal processes in Pakistani Halys sindillus Memon, Meier et Manan, 2006. Considering the data summarised above, there is no wonder that the authors providing comprehensive treatments of Halyini (WALL 2004, MEMON et al. 2011) found no clear-cut synapomorphies for defining the tribe. Within the Madagascan fauna, Carduelicoris gen.nov. fits to Halyini in the shape of the head, sharing with typical forms of the tribe the elongate head with mandibular plates not particularly enlarged and foliaceous (in contrast with Triplatygini and Rolstoniellini), but usually bearing prominent denticles or spines (in contrast with the unarmed head of Amyntorini) (CACHAN 1952). The prominent humeral angles of pronotum were mentioned as characteristic for Halyini by GROSS (1976); however, none of the Madagascan genera of Halyini treated by CACHAN (1952) (nor any other Halyini genus known to me) has the humeral angles of the pronotum reaching so far anteriad and being apically lobate or spinous. Such shape of humeral angles is otherwise characteristic for the endemic Madagascan tribe Triplatygini (see CACHAN 1952; KMENT 2008, 2012), the Oriental genus Rolstoniellus Rider, 1997 (= Compastes Stål, 1867, Rolstoniellini) (DISTANT 1902, YANG 1935), or the Chinese Brachymna humerata Chen, 1989 (Sephelini) (CHEN 1989), but I consider these similarities as parallelisms. Considering the genital characters, both male (strongly sclerotised phallotheca, number of conjunctival processes, shape of ventral conjunctival processes, and especially the gonopore opening between the ventral conjunctival processes and the vesica not protruding beyond them (see GROSS 1976)) and female genitalia (shape and number of processes of the apical receptacle of spermatheca, shape of the sclerotised rod) of Carduelicoris gen.nov. seem to fit into the variability of genitalic structures known in this group, except for the presence of the dorsal processes 1) in pygophore which seems quite exceptional within the Halyini, since they are apparently missing in most members of the tribe (e.g., GROSS 1972, 1976; GHAURI 1982; ABBASI 1986; LINNAVUORI 1982; WALL 2007; MEMON & AHMAD 2008; TSAI & RÉDEI 2009; FAN 2011; MEMON et al. 2011; FAN & LIU 2013). Dorsal processes are sclerotized appendages of the intersegmental membrane between

1) This structure is termed variously in the literature: superior lateral processes (SHARP 1890), genital plate (e.g. BAKER 1931; MCDONALD 1966, 1986), dorsal process (LESTON 1954, DAVIDOVÁ-VILÍMOVÁ & MCPHERSON 1992), processus supérieurs (DUPUIS 1951, 1970), superior process (FERRARI et al. 2010), pseudoclasper (MCDONALD 1974, 1976, 1982; FAN 2011), pygophoral plate (ROLSTON & MCDONALD 1979), parandrium (GROSS 1975, 1976), dorsal plate (LINNAVUORI 1982, KMENT & JINDRA 2009), or genital cup process (BARCELLOS & GRAZIA 2003).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 427 P. K MENT segments IX and X (LESTON 1954) supposed to give support to parameres (LINNAVUORI 1982). These structures occur in number of genera of Asopinae, Edessinae, and Pentatominae (e.g. in the tribes Aeliini, Aeptini, Antestiini, Bathycoeliini, Cappaeini, Carpocorini, Caystrini, Diemeniini, Euryaspidini, Eysarcorini, Myrocheini, Nezarini, Pentatomini, Procleticini, Sephelini, and Strachiini), and most probably evolved several times independently in different evolutionary lineages (see e.g. BAKER 1931; MCDONALD 1966, 1974, 1976, 1982, 1986, 1995; MCDONALD & EDWARDS 1978; GROSS 1975, 1976; ROLSTON & MCDONALD 1979; LINNAVUORI 1982; DAVIDOVÁ-VILÍMOVÁ & MCPHERSON 1992; BARCELLOS & GRAZIA 2003; KMENT & JINDRA 2009; FERRARI et al. 2010; FAN 2011). Carduelicoris gen.nov. differs from Triplatyx Horváth, 1904 and Anoano Cachan, 1952 of Triplatygini also in the absence of the triangulin; both latter genera have the triangulin well-developed (KMENT 2008, 2012). Considering all the characters, Carduelicoris gen.nov. is placed here within Halyini, and the characters shared with Triplatygini are considered either symplesiomorphic (short spout-shaped peritreme, abdominal base without conical tubercle) or convergent (prominent humeral angles of pronotum).

Diversity of Halyini in the Afrotropical Region and Madagascar Halyini (including Memmiini, see LINNAVUORI 1982 for the synonymy) is one of the diverse and species-rich tribes of Pentatominae, including 84 genera and about 400 species (LINNAVUORI 1982, WALL 2007, MEMON & AHMAD 2008, FAN & LIU 2013, RIDER 2013) with most species distributed in the Oriental and Australian Regions, also with considerable diversity in the Afrotropical Region, Madagascar, the southern areas of the Palaearctic Region, and with two endemic genera in North and Central America (CACHAN 1952; GROSS 1972, 1976; LINNAVUORI 1982; ABBASI 1986; LARIVIÈRE 1992, 1994; CASSIS & GROSS 2002; RIDER 2006; WALL 2007; MEMON et al. 2011; etc.). The continental African fauna of Halyini was partly reviewed by LINNAVUORI (1982), who divided the African genera into two groups: (1) Atelocera group limited to the Afrotropical Region and extending to Yemen and Madagascar, comprising the genera Atelocera Laporte, 1833, Memmia Stål, 1865, Mezessea Linnavuori, 1982, Pseudatelus Linnavuori, 1982, and Scribonia Stål, 1865, and (2) Halys group, considered by LINNAVUORI (1982) as of Oriental origin, including the genera Halys Fabricius, 1803, Erthesina Spinola, 1837 (both shared with the Oriental Region), Coenomorpha Dallas, 1851 (widely distributed in tropical Africa), Eurus Dallas, 1851, Orthoschizops Spinola, 1850 (both endemic to South Africa), and Anolcus Bergroth, 1893 (endemic to East Africa). Compared with continental Africa, the Madagascan fauna of Halyini is more diversified and highly endemic, with eleven of the thirteen genera and apparently all 40 described species restricted exclusively to the island: Abadia Cachan, 1952 (1 species), Acuticeps Cachan, 1952 (1 species), Artiazontes Distant, 1881 (1 species), Bourginia Cachan, 1952 (1 species), Carduelicoris sp.nov. (1 species), Dalpada Amyot et Serville, 1843 (16 species), Halys (1 species), Memmia (6 species), Nesagaeus Bergroth, 1906 (1 species), Nesobius Bergroth, 1906 (2 species), Solenogaster Reuter, 1867 (1 species), Vadonidea Cachan, 1952 (2 species), and Zaplutus Bergroth, 1893 (6 species) (CACHAN 1952, this

428 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Carduelicoris stehliki gen. et sp.nov. of Pentatomidae paper). However, the Madagascan Halyini remain poorly known, especially in respect of the detailed structures of the male and female genitalia. For example, the Madagascan species currently accommodated in Dalpada seem heterogenous even for a superficial look, and apparently they represent at least three separate genera, neither of them being congeneric with the Oriental Dalpada aspersa Amyot et Serville, 1843, the type species of Dalpada; the same may be true also for the Madagascan and Afrotropical Halys species. It is also probable that the endemic Madagascar tribe Nealeriini, including the genera Nealeria Bergroth, 1893 (2 species) and Paraleria Reuter, 1887 (1 species), both of Halyini-like habitus and apparently differing from the latter tribe only by the presence of 2- segmented tarsi, also belongs to the same evolutionary lineage and is therefore merely a subgroup of a more broadly defined Halyini (see also WALL 2004). In the future, a thorough revision is needed to clarify the identity of the Madagascan genus-group taxa currently placed to Fig. 26. Distribution map of Carduelicoris stehliki gen. Halyini and their phylogenetic and et sp.nov. (type locality marked with T). zoogeographic relationships with other Afrotropical and Oriental taxa.

Acknowledgements I am obliged to Dominique Pluot-Sigwalt and Eric Guilbert (MNHN), and Mick Webb (BMNH) for loans of the specimens under their care and kind assistance during my stays in Paris and London, and to Dávid Rédei (Nankai University, Tianjin, China) and David Rider (North Dakota University, Fargo, USA) for many valuable comments on the manuscript. This research received support from the SYNTHESYS Project http://www.synthesys.info/ which is financed by European Community Research Infrastructure Action under FP7 Integrating Activity Programme. The preparation of this manuscript received financial support from the Ministry of Culture of the Czech Republic to the National Museum, Prague (DKRVO MK-S 760/2013/12 OVV).

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References

ABBASI Q. L. 1986: Morpho-taxonomic studies of the family Pentatomidae Leach, 1815 (Heteroptera: Pentatomomorpha) of South Asia (Pakistan, Azad Kashmir and Bangladesh) with reference to the phylogeny of the group. Supplement of Pakistan Journal of Entomology Karachi 5: 1–279. AHMAD I. 1995: A review of pentatomine legume bug genus Piezodorus Fieber (Hemiptera: Pentatomidae: Pentatominae) with its cladistic analysis. Proceedings of the Pakistan Congress of Zoology 15: 329–358. ARNOLD K. 2011: Jostenicoris Arnold, n. gen., eine neue Heteropteren-Gattung aus der Äthiopischen Region (Insecta: Hemiptera: Heteroptera: Pentatomidae: Strachiini). Edessana 1: 15–18. BAKER A. D. 1931: A study of the male genitalia of Canadian species of Pentatomidae. Canadian Journal of Reasearch 4(2): 148–179, 4(3): 181–220 + pls. I–II. BARCELLOS A. & GRAZIA J. 2003: Cladistic analysis and biogeography of Brachystethus Laporte (Heteroptera, Pentatomidae, Edessinae). Zootaxa 256: 1–14. CACHAN P. 1952: Les Pentatomidae de Madagascar (Hemipteres Heteropteres [sic!]). Mémoires de l’Institut Scientifique de Madagascar, Série E 1: 231–462. CASSIS G. & GROSS G. F. 2002: Hemiptera: Heteroptera (Pentatomomorpha). In: HOUSTON W. W. K. & WELLS A. (eds.): Zoological Catalogue of Australia. Vol. 27.3B. CSIRO, Melbourne, xiv + 737 pp. CHEN ZH.-Y. 1989: [Two new species of Pentatominae from Guangdong, China (Hemiptera: Pentatomidae)]. Acta Entomologica Sinica 32: 477–479 (in Chinese, English summary). COUILLOUD R. 1989: Hétéroptères déprédateurs du cotonnier en Afrique et à Madagascar (Pyrrhocoridae, Pentatomidae, Coreidae, , , Lygaeidae). Coton et Fibre Tropicales 44: 185–227. DAVIDOVÁ-VILÍMOVÁ J. & MCPHERSON J. E. 1992: Pygophores of selected species of Pentatomoidea (Heteroptera) from Illinois. Acta Universitatis Carolinae Biologica 36: 143–183. DAY G. M. 1965: Revision of Acrosternum Auctt. nec Fieber from Madagascar. Annals and Magazine of Natural History, Series 13 7 (1964): 559–565. DERJANSCHI V. & P ÉRICART J. 2006: Hémiptères Pentatomoidea Euro-Méditerranéens. Volume I. Faune de France. Vol. 90. Fédération Française des Sociétés de Sciences Naturelles, Paris, 494 pp + 16 pls. DISTANT W. L. 1902: Rhynchota. – Vol. I. (Heteroptera). In: BLANFORD W. T. (ed.): The fauna of British India including Ceylon and Burma. Taylor and Francis, London, xxxviii + 438 pp. DUPUIS C. 1951: Les espèces françaises du genre Eurydema Laporte (Hemiptera, Pentatomoidea, subfam. Pentatominae). Révision systématique avec une introduction à l’étude morphologique des organes génitaux externes des Pentatomoidea. Annales de la Société Entomologique de France 118: 1–28. DUPUIS C. 1970: Heteroptera. Pp. 190–209. In: TUXEN S. L. (ed.): Taxonomist’s glossary of genitalia in insects, 2nd edition. Munksgaard, Copenhagen, 359 pp. FAN ZH.-H. 2011: [Study on the taxonomy of Pentatominae from China (Hemiptera: Heteroptera: Pentatomidae)]. Unpublished Ph.D. Thesis. Nankai University, Tianjin, ix + 614 pp (in Chinese, English abstract). FAN ZH.-H. & LIU G.-Q. 2013: The genus Cahara Ghauri, 1978 of China (Hemiptera, Heteroptera, Pentatomidae, Halyini) with descriptions of two new species. ZooKeys 319: 37–50. FERRARI A., SCHWERTNER C. F. & GRAZIA J. 2010: Review, cladistic analysis and biogeography of Nezara Amyot & Serville (Hemiptera: Pentatomidae). Zootaxa 2424: 1–41. GHAURI M. S. K. 1982: New genera and new species of Halyini, mainly from South India (Heteroptera, Pentatomidae, Pentatominae). Reichenbachia 20: 1–24. GÖLLNER-SCHEIDING U. 1999. Die Gattung Phricodus Spinola, 1840 (Insecta: Heteroptera: Pentatomidae). Entomologische Abhandlungen Staatliches Museum für Tierkunde Dresden 58: 149– 163. GREATHEAD D. J. 1968: On the taxonomy of Antestiopsis spp. (Hem., Pentatomidae) of Madagascar, with notes on their biology. Bulletin of Entomological Research 59: 307–315. GROSS G. F. 1972: A revision of the species of Australian and New Guinea shield bugs formerly placed in the genera Poecilometis Dallas and Eumecopus Dallas (Heteroptera: Pentatomidae) with descriptions of new species and selection of lectotypes. Australian Journal of Zoology, Supplement 15: 1–192. GROSS G. F. 1975: Plant feeding and other bugs (Hemiptera) of South Australia – Heteroptera. Part I. A. B. James, Adelaide, South Australia, pp. 1–250. GROSS G. F. 1976: Plant feeding and other bugs (Hemiptera) of South Australia – Heteroptera. Part II. A. B. James, Adelaide, South Australia, pp. 251–501.

430 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Carduelicoris stehliki gen. et sp.nov. of Pentatomidae

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE (ICZN) 1999: International Code of Zoological Nomenclature. Fourth edition. International Trust for Zoological Nomenclature, London, 306 pp. KMENT P. 2008: A revision of the endemic Madagascan genus Triplatyx (Hemiptera: Heteroptera: Pentatomidae). Acta Entomologica Musei Nationalis Pragae 48: 543–582. KMENT P. 2011: Triplatyx ribesi sp. nov., a new species of Pentatomidae (Hemiptera: Heteroptera) from Madagascar. Heteropterus Revista de Entomología 11: 279–286. KMENT P. 2012: Redescription of the Madagascan endemic genus Anoano with a new synonymy (Hemiptera: Heteroptera: Pentatomidae). Acta Entomologica Musei Nationalis Pragae 52: 371–382. KMENT P. & J INDRA Z. 2009: A revision of Tripanda and Tenerva (Hemiptera: Heteroptera: Pentatomidae: Pentatominae). Zootaxa 1978: 1–47. KMENT P. & V ILÍMOVÁ J. 2010: Thoracic scent efferent system of Pentatomoidea (Hemiptera: Heteroptera): a review of terminology. Zootaxa 2706: 1–77. LARIVIÈRE M.-C. 1992: Description of Parabrochymena, new genus, and redefinition and review of Brochymena Amyot and Audinet-Serville (Hemiptera: Pentatomidae), with considerations on natural history, chorological affinities, and evolutionary relationships. Memoirs of the Entomological Society of Canada 163: 1–75. LARIVIÈRE M.-C. 1994: Parabrochymena Larivière (Hemiptera: Pentatomidae): Systematics, natural history, chorological affinities, and evolutionary relationships, with a biogeographic analysis of Parabrochymena and Brochymena Amyot and Audinet-Serville. Canadian Entomologist 126: 1193–1250. LESTON D. 1953a: Notes on the Ethiopian Pentatomoidea (Hem): vii* A review of Gynenica Dallas 1851. Revue de Zoologie et de Botanique Africaine 48: 179–195. LESTON D. 1953b: Notes on the Ethiopian Pentatomoidea (Hem.): XIII. On a waxy secretion by Halyini (Pentatomidae). Entomologist’s Monthly Magazine 89: 147. LESTON D. 1954: Notes on the Ethiopian Pentatomoidea (Hem.). XII. On some specimens from Southern Rhodesia, with an investigation of certain features in the morphology of Afrius figuratus (Germar) and remarks upon the male genitalia of Amyotinae. Occasional Papers of the National Museum of Southern Rhodesia 19: 678–686. LINNAVUORI R. 1973: Studies on African Heteroptera. Arquivos do Museu Bocage, 2.a Série 4(2): 29–69. LINNAVUORI R. 1975: Hemiptera Heteroptera of the Sudan with remarks on some species of the adjacent countries. Part 5. Pentatomidae. Boletim da Sociedade Portuguesa de Ciências Naturais, Série 2 15: 5–128. LINNAVUORI R. 1982: Pentatomidae and (Heteroptera) of Nigeria and the Ivory Coast, with remarks on species of the adjacent countries in West and Central Africa. Acta Zoologica Fennica 163: 1–176. MCDONALD F. J. D. 1966: The genitalia of North American Pentatomoidea (Hemiptera: Heteroptera). Quaestiones Entomologicae 2: 7–150. MCDONALD F. J. D. 1974: Revision of the genus Holcostethus in North America (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 82: 245–258. MCDONALD F. J. D. 1976: Revision of the genus Trichopepla (Hemiptera:Pentatomidae) in N. America. Journal of the New York Entomological Society 84: 9–22. MCDONALD F. J. D. 1982: Description of the male genitalia of Holcostethus hirtus (Van Duzee) with a revised key to North American species (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 90: 5–7. MCDONALD F. J. D. 1986: Revision of Cosmopepla Stål (Hemiptera: Pentatomidae). Journal of the New York Entomological Society 94: 1–15. MCDONALD F. J. D. 1995: Revision of Hypogomphus Spinola in Australia (Hemiptera: Pentatomidae). Journal of the Australian Entomological Society 34: 169–176. MCDONALD F. J. D. & EDWARDS P. B. 1978: Revision of the genus Oncocoris Mayr (Hemiptera: Pentatomidae). Australian Journal of Zoology, Supplementary Series 62: 1–53. MEMON N. & AHMAD I. 2008: Description of Mustha izmirensis, new species (Heteroptera: Pentatomidae: Halyini) from Bornova, Izmir, Turkey with key to its world species. Pakistan Journal of Zoology 40: 435–439. MEMON N. & AHMAD I. 2009: A revision of Halyine stink bug genus Sarju Ghauri (Hemiptera: Pentatomidae: Halyini) and its cladistic analysis. Pakistan Journal of Zoology 41: 399–411.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 431 P. K MENT

MEMON N., GILBERT F. & AHMAD I. 2011: Phylogeny of the South Asian Halyine stink bugs (Hemiptera: Pentatomidae: Halyini) based on morphological characters. Annals of the Entomological Society of America 104: 1149–1169. MEMON N., MEIER R., MANAN A. & SU K. F.-Y. 2006: On the use of DNAsequences for determining the species limits of a polymorphic new species in the stink bug genus Halys (Heteroptera: Pentatomidae) from Pakistan. Systematic Entomology 31: 703–710. ORIAN A. J. E. 1965: A new genus of Pentatomidae from Africa, Madagascar and Mauritius (Hemiptera). Proceedings of the Royal Entomological Society of London (B) 34: 25–30. RIDER D. A. 1998a: Nomenclatural changes in the Pentatomoidea (Hemiptera-Heteroptera: Cydnidae, Pentatomidae). II. Species level changes. Proceedings of the Entomological Society of Washington 100: 449–457. RIDER D. A. 1998b: Nomenclatural changes in the Pentatomoidea (Hemiptera-Heteroptera: Pentatomidae, ). III. Generic level changes. Proceedings of the Entomological Society of Washington 100: 504–510. RIDER D. A. 2006: Family Pentatomidae. Pp. 233–402. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 5. The Netherlands Entomological Society, Amsterdam, xiii + 550 pp. RIDER D. A. 2007: Cachaniellus, a replacement name for the preoccupied genus Cachanocoris (Hemiptera: Heteroptera: Pentatomidae). Proceedings of the Entomological Society of Washington 109: 261. RIDER D. A. 2013: Pentatomoidea Home page. Available online at http://www.ndsu.nodak.edu/ndsu/rider/ Pentatomoidea/ (accessed on August 15, 2013). ROLSTON L. H. & MCDONALD F. J. D. 1979: Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera). Journal of the New York Entomological Society 87: 189–207. SCHAEFER C. W. 1977: Genital capsule of the trichophoran male (Hemiptera: Heteroptera: Geocorisae). International Journal of Insect Morphology & Embryology 6: 277–301. SCHOUTEDEN H. 1954: Deux Pentatomides nouveaux de Madagascar. Revue de Zoologie et de Botanique Africaine 49: 6–8. SHARP D. 1890: On the structure of the terminal segment in some male Hemiptera. Transactions of the Entomological Society of London 1890(3): 399–430, pls. 12–14. ŠTYS P. & EXNEROVÁ A. 2003: Ventral surface of forewings in Pentatomidae: Halyini (Heteroptera), an overlooked site of fulguration. Acta Universitatis Carolinae Biologica 47: 275–284. TSAI J.-F. & RÉDEI D. 2009: The identity of shield bugs described by Francis Walker from Taiwan (Hemiptera: Heteroptera: Pentatomidae). Zootaxa 2152: 43–54. TSAI J.-F., RÉDEI D., YEH G.-F. & YANG M.-M. 2011: Jewel bugs of Taiwan (Heteroptera: ). National Chung Hsing University, Taichung, 309 pp. WALL M. A. 2004: Phylogenetic relationships among Halyini (Pentatomidae: Pentatominae) genera based on morphology, with emphasis on the taxonomy and morphology of the Solomonius-group. Unpublished Ph.D. dissertation. University of Connecticut, Storrs. [Not seen, fide MEMON et al. (2011)]. WALL M. A. 2007: A revision of the Solomonius-group of the stinkbug tribe Halyini (Hemiptera: Pentatomidae: Pentatominae). Zootaxa 1539: 1–84. YANG W.-I. 1935: Description on three new species of gen. Compastes (Pentatomidae, Hem.-Het.). Bulletin of the Fan Memorial Institute of Biology, Zoology 6(3): 98–102.

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Behaviour of the central European Acanthosomatidae (Hemiptera: Heteroptera: Pentatomoidea) during oviposition and parental care

JANA HANELOVÁ & JITKA VILÍMOVÁ Charles University, Faculty of Science, Department of Zoology, Vinièná 7, CZ-128 44, Praha 2, Czech Republic; e-mail: [email protected], [email protected]

HANELOVÁ J. & VILÍMOVÁ J. 2013: Behaviour of the central European Acanthosomatidae (Hemiptera: Heteroptera: Pentatomoidea) during oviposition and parental care. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 433–457. – Six central European Acanthosomatidae: Acanthosomatinae species were studied with particular reference to the behaviour of females during and after oviposition. Acanthosoma haemorrhoidale (Linnaeus, 1758), Cyphostethus tristriatus (Fabricius, 1758), and Elasmostethus interstinctus (Linnaeus, 1758) females do not exhibit maternal care. Eggs are deposited in many small batches as a strategic adaptation against predators and parasitoids. In contrast, Elasmucha ferrugata (Fabricius, 1787), Elasmucha fieberi Jakovlev, 1864, and Elasmucha grisea (Linnaeus, 1758) do exhibit maternal care. The defensive behaviour of the Elasmucha species is almost identical. Reactions to a potential aggressor graduate from fine to strong intensity. If the female leaves her eggs while feeding, she cannot distinguish her own eggs upon return, only the eggs of her own species.

Keywords. Pentatomomorpha, Acanthosomatidae, true bugs, Acanthosoma haemorrhoidale, Cyphostethus tristriatus, Elasmostethus interstinctus, Elasmucha ferrugata, Elasmucha fieberi, Elasmucha grisea, behaviour, maternal care

Introduction Parental care of offspring is a subsocial behaviour occurring quite often within the Arthropoda, including . Members of 13 insect orders tend and defend their immature stages, including eggs (TALLAMY & WOOD 1986, TALLAMY 2001). Heteroptera have been the focus of numerous studies, because some of them show maternal care, also – if rarely – paternal care (e.g. SMITH 1997, summary; TALLAMY & SCHAEFER 1997). Parental care is distributed across several recent heteropteran taxa. In addition to Belostomatidae, paternal care has been mentioned in Coreidae and (e.g. KAITALA et al. 2001, TALLAMY 2001), and maternal care is known in a total of 15 families (HANELOVÁ 2005). The generally accepted hypothesis is that parental care is an advanced mode of behaviour. However, TALLAMY & SCHAEFER (1997) have suggested quite the opposite: that the care may be a surviving ancestral pattern of reproductive behaviour. Parental care provides effective protection of the offspring against the predators, parasites, and parasitoids that regularly increase mortality in the species. Important comprehensive studies of parental care in the Heteroptera include, for example, COBBEN (1968), RIDLEY (1978), TALLAMY & WOOD (1986), TALLAMY & SCHAEFER (1997), and TALLAMY (2000, 2001). TALLAMY & WOOD (1986) distinguished three basic patterns of

433 J. HANELOVÁ & J. VILÍMOVÁ care behaviour: (i) physical protection of egg batches and young larvae by an adult (many studies); (ii) protection of food resources or providing food for larvae in Cydnidae and (e.g. SITES & MCPHERSON 1982, FILIPPI et al. 2000, NOMAKUCHI et al. 1998); (iii) female feeding the larvae in (e.g. BEQUAERT 1935, TSUKAMOTO & TOJO 1992, GUILBERT 2003). Maternal care occurs within the heteropteran superfamily Pentatomoidea, particularly in members of the families Acanthosomatidae (see below), Cydnidae (e.g. KIGHT 1996, FILIPPI-TSUKAMOTO et al. 1995), and Tessaratomidae (e.g., GOGALA et al. 1998, MONTEITH 2006). This subsocial pattern of behaviour in the Acanthosomatidae has been known for 250 years or more, first recorded in the 18th century. It was first studied in detail in a few European species, and later in Japanese ones (TALLAMY 1999). Maternal care is known in eleven species from four acanthosomatid genera, Anaxandra Stål, 1876, Elasmucha Stål, 1864, Sastragala Amyot & Serville, 1843, and Sinopla Signoret, 1863 (Table 1). Most species with parental care (seven) are of the genus Elasmucha. All three central European Elasmucha species (E. ferrugata (Fabricius, 1787), E. fieberi Jakovlev, 1864, and E. grisea (Linnaeus, 1758)) exhibit parental care. The first published note on parental care in Acanthosomatidae was an observation on E. grisea (MODEER 1764, after TALLAMY 1999), also the most studied acanthosomatid species in recent years (e.g. MELBER et al. 1980, KAITALA & MAPPES 1997; Table 1). In contrast, E. fieberi and E. ferrugata have been studied far less (e.g. KAITALA & MAPPES 1997). No detailed recent data, except a study by FISCHER (2006), mention possible care in other central European acanthosomatids. The behaviour of an Elasmucha grisea female while she cared for her brood was studied in detail, including experiments designed to elicit defence reactions (MELBER & SCHMIDT 1975a, MAPPES & KAITALA 1994). MELBER & SCHMIDT (1975a) and later MELBER et al. (1980) classified defence behaviour into five stages. More studies have been dedicated to defence behaviour in Elasmucha dorsalis (Jakovlev, 1876) and Elasmucha signoreti Scott, 1874 (e.g. KUDO et al. 1989, KUDO & NAKAHIRA 1993). A detailed account of the behaviour of the female E. ferrugata in the course of parental care is the central aim of this paper. However, other central European Acanthosomatidae were also observed in terms of care for offspring including eggs.

Material and methods Material. All the Acanthosomatidae material for this study was collected in central Bohemia, in the wider surroundings of Vlašim (map grid 6255, after PRUNER & MÍKA 1996) (see HANEL & HANELOVÁ 2007). Two species, Elasmucha ferrugata (Fabricius, 1787) and Elasmucha grisea (Linnaeus, 1758), were observed in detail and used in experiments. Four species, Acanthosoma haemorrhoidale (Linnaeus, 1758), Cyphostethus tristriatus (Fabricius, 1758), Elasmostethus interstinctus (Linnaeus, 1758) and Elasmucha fieberi Jakovlev, 1864, were studied only partially, for oviposition mode and parental care (Table 2). The numbers of specimens of particular Acanthosomatidae species are summarized in Table 2.

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Breeding. Acanthosomatids were bred in small plastic containers or in larger cylindrical containers made of gauze, both covered with very fine netting (Figs 1, 2). The host plants (Table 3) were placed in glass containers with water. The containers were kept in normal room conditions (approximate temperature 18o–30oC). Observations and experiments. Records of the experiments were kept with a Nikon D 70 camera, and the plates were prepared with the aid of Adobe PhotoShop software. Direct observation of the behaviour of the bugs, either in their breeding containers or natural conditions, was carried out several times a day, on average five. Experiments were performed in subject bugs in breeding containers as well as in natural conditions. A range of animals and equipment was used to elicit defensive reactions: Chelicerata: Acarina: Ixodes sp. (Ixodidae), body length 2 mm; Hexapoda: Coleoptera: Chrysomela populi Linnaeus, 1758 (Chrysomelidae), body length 10 mm; Phyllopertha horticola (Linnaeus, 1758) (Scarabaeidae), body length 10 mm; Rhagonycha fulva (Scopoli, 1763) (Cantharidae), body length 9 mm; Hymenoptera: Formica polyctena Förster, 1850 (Formicidae), body length 7 mm; Diptera: Asilidae gen. sp., body length 17 mm; other stages of the same Elasmucha species, body length 7 mm; other Elasmucha species, body length 7 mm; tips of soft tweezers; and a magnifying glass 13 cm in diameter. The species represented possible natural enemies that Elasmucha species might encounter in nature. MELBER et al. (1980) used similar insect taxa in their experiments with E. grisea. The experiment of intensity of care involved eleven females of E. ferrugata and their hatched eggs; they took care of eggs and also of young larvae. Within a period of 120 minutes every day, the female was intruded upon from the side with soft tweezers (15 cm long, 2 mm wide at the tip) every five minutes (Fig. 3). The females reacted in repeatedly recognisable fashion. There was a body shift that turned the female to face the threat every time. After reaction and retraction of threat, the female returned to her default position, i.e. parallel to the surface of the leaf with the egg batch. Slight intensity reactions • Movements of the antennae – quick changes of antennal position, such as straightening, lifting, and jerking. Medium intensity reactions • Covering the egg batch with the female’s body, partially or completely. • Body tilting, facing the threat. Strong intensity reactions • Body jerking – quick side-to-side movements of the body (= short changes of horizontal position of entire body). • Body swinging – slower changes of body position from left to right and back again. • Wing-fanning – short, quick, spinning movements of the partly-outspread wings, both pairs.

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Figs 1–6. 1 – small breeding container; 2 – large breeding container; 3 – Elasmucha ferrugata (Fabricius, 1787) female intruded upon with soft tweezers; 4 – Elasmucha ferrugata male attempting to copulate with female over an egg batch; 5 – Elasmucha grisea (Linnaeus, 1758) female with clutch on the dorsal surface of a birch leaf; 6 – Elasmucha fieberi Jakovlev, 1864 female with clutch on a birch leaf.

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Figs 7–12. 7 – egg batch of Elasmostethus interstinctus (Linnaeus, 1758); 8 – ovipositing female of Cyphostethus tristriatus (Fabricius, 1758); 9 – egg batch of Cyphostethus tristriatus on Juniperus communis L. berry; 10 – Elasmostethus interstinctus sucking on conspecific eggs; 11 – larvae of Cyphostethus tristriatus aggregated on Juniperus communis berry; 12 – first instar larvae of Acanthosoma haemorrhoidale (Linnaeus, 1758) on a rowan leaf.

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Results If no information is available for a given species, it is not mentioned. The numbers of specimens used in the experiments are listed in Table 2.

Copulation Acanthosoma haemorrhoidale, Cyphostethus tristriatus, Elasmostethus interstinctus, Elasmucha ferrugata, and E. grisea: Pairs of all species in copula assumed a tandem position. In two pairs of E. ferrugata, the male tried to copulate with a female over an egg batch. The male was on the female’s back, adequate for the first stage of copulation, but the pairing was not successfully completed (Fig. 4).

Oviposition and size of egg batch (Table 4) Females lay their eggs only after feeding on host plants with fruit, during the night in E. ferrugata and E. grisea but during the day in Acanthosoma haemorrhoidale and Cyphostethus tristriatus. Females of Elasmucha ferrugata (n = 25) laid 33–39 eggs, without exception on the ventral surface of host plant leaves, in containers as well as in natural conditions. Females of Elasmucha grisea (n = 20) laid 35–45 eggs on either surface of leaves; of 70 batches, 30 were on a dorsal surface, 40 on a ventral surface (Fig. 5). A female of Elasmucha fieberi laid 34 whitish eggs (Fig. 6). The eggs of all Elasmucha species are whitish. Females of Elasmostethus interstinctus laid smaller egg batches, 4–30 (Fig. 7), on either surface of the host plant leaves. The eggs are green. Cyphostethus tristriatus deposited similarly small egg batches, with 3–20 eggs in each eggs in each batch (Figs 8–9), on needles and berries of the host plant. One female of C. tristriatus did not deposit an egg batch all at once, but did so by stages. The eggs are pale green. Females of E. ferrugata can deposit a second egg batch (32 % = 16 observed specimens) and eventually a third batch (8 % = 4 specimens), particularly when the first egg batch or larvae have been destroyed. This species occurred and oviposited first on bilberry (Vaccinium myrtillus), often moving on to fly honeysuckle (Lonicera xylosteum), where it (potentially) deposited a second egg batch. Four females in breeding containers continued their care even after all the larvae of the first instar had died. They stood over them for 4–6 days; this behaviour was not observed in E. grisea. Elasmostethus interstinctus and Cyphostethus tristriatus deposited larger numbers of egg batches, for which they showed no inclination to care at all: the females simply left the Fig. 13. Intensity of defence of eggs and larvae in egg batch immediately after Elasmucha ferrugata (Fabricius, 1787) females. oviposition. Cannibalism, feeding on

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Figs 14–18. 14 – Elasmucha ferrugata (Fabricius, 1787) female responds to the presence of a conspecific male; 15 – E. ferrugata female responds to the presence of Chrysomela populi (Linnaeus, 1758); 16 – E. ferrugata female feeding on the host plant, away from eggs; 17–18 – E. ferrugata female adopting conspecific first-instar larvae.

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Figs 19–22. Female of Elasmucha grisea (Linnaeus, 1758). 19 – female shows no interest in larvae of Elasmostethus interstinctus (Linnaeus, 1758); 20 – two females with egg batches on one leaf; 21 – two females with hatched larvae on one leaf; the female on the right wing-jerking at the other; 22 – the stronger female adopts the larvae of the weaker one.

the eggs of their own species, was observed in E. interstinctus (Fig. 10). Acanthosoma haemorrhoidale laid 2–28 whitish eggs, always located on the ventral side of a rowan (Sorbus aucuparia) leaf. This species also lays her eggs in several batches, as do other species that lack care (HANEL & HANELOVÁ 2008).

Parental care Acanthosoma haemorrhoidale, Cyphostethus tristriatus, and Elasmostethus interstinctus. Females of all these species left the egg batches immediately after oviposition and never returned. Parental care does not appear to occur in these species. First and second instars aggregated on, or close to, food resources (Figs 11, 12). 440 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Behaviour of Acanthosomatidae during oviposition and parental care

Elasmucha ferrugata, E. fieberi, and E. grisea. Females of all three species exhibit maternal care of the egg batch and young larvae. Behaviour was studied in detail only for E. ferrugata and E. grisea. The female stood over the egg batch throughout egg development. First instars barely moved after hatching, keeping in a compact group with the female, just sucking on their empty egg shells. The second instar moved towards food – berries or catkins – with the female in close attendance. She checked the larvae constantly and managed them with touches of her antennae. Elasmucha ferrugata cared for the larvae until the second instar. Care was the most intensive for the first instar. The female did not stay with the second instar all the time, but remained close to where it was feeding. The female returned to the larvae for the night. Females of E. grisea cared for the larvae until the third instar. The female stood over the eggs and first instar aggregated on empty egg shells. The second instar was similarly gregarious and the female remained with it. During the third instar, larvae moved towards food and back to the leaf and the female guarded them. Larvae split into smaller groups and dispersed at the end of the third instar, at which point the female left them. Behaviour of the females with eggs and larvae. The female in repose over the egg batch keeps her antennae directed backwards along the body. If stimulated, she straightens them and directs them forwards. Other types of reaction follow, not necessarily in the same order every time (Table 5). A hundred responses to “attack” with soft tweezers were recorded in 20 E. ferrugata females. Defensive behaviour began when the tweezers were about 10 cm from them. A total of 12 combinations of reaction were recognized. The most frequent sequence (37 %) was: straightening the antennae – body tilting against the attack – body jerking. All possible reactions were recorded in 6 % of cases. Body jerking and wing-fanning occurred as strongly intensive reactions; they might be repeated several times. Similar experiments were carried out on an E. grisea female; these served only as a control, since the behaviour of that species is well known. Ten E. ferrugata females were “attacked” by an approaching magnifying glass. All of them responded with combinations of antenna-straightening – body-jerking – wing- fanning. They also reacted to the movements of a relatively huge intruder, i.e. a person, usually by pressing themselves more strongly against the egg batch when the threat was about one metre away. The females defended larvae at a markedly higher level of intensity than they showed for the eggs (Fig. 13). In an experiment that lasted two hours, significantly higher frequencies of antennal movement, covering of the egg batch with the body, body- jerking, and wing-fanning were recorded when the larvae were being defended in comparison with the eggs. Experimental attack by other arthropods. The females of E. ferrugata reacted to the presence of other arthropods with: slight intensity of reaction towards Ixodes sp.; and medium intensity to other specimens of E. ferrugata (Fig. 14), Chrysomela populi (Fig. 15), Phyllopertha horticola, Rhagonycha fulva, and Asilidae gen., sp. An ant, Formica polyctena, attacked an E. ferrugata female with eggs. She promptly left the clutch and only returned to it after the ant had been removed. Similarly, E. grisea females reacted

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 441 J. HANELOVÁ & J. VILÍMOVÁ only slightly to the presence of a conspecific male but strongly to Rhagonycha fulva. A female of E. grisea with eggs actively attacked larvae of E. ferrugata occurring close to her. Female feeding during care. Females of both Elasmucha species were observed to feed during their care of egg batches. Five females of E. ferrugata left the egg batch for various time periods, from 30 minutes to six hours, and sucked from fresh parts of the host plant (Fig. 16). Two of these females were minding a second egg batch. The females of E. grisea exhibited identical behaviour. Egg batch adoption. In the course of the experiments, all 25 E. ferrugata females adopted a conspecific egg batch that was not their own, for various periods of time from 39 to133 minutes. An E. ferrugata female bred in the container took over care of whatever batch she found first, regardless of any parental relationship. On finding a clutch, the female checked it with her antennae and rostrum for 9–16 minutes, and then took up position over it. Larvae could also be adopted; an E. ferrugata female took over the larvae of other females (Figs 17, 18), whether actively or if new larvae became mixed in with her own. Egg batches of different but congeneric species, i.e. E. grisea, were not adopted by E. ferrugata. The latter touched the batch with the antennae and rostrum for a couple of minutes, then left. Females of E. grisea exhibited identical behaviour in terms of adoption of other clutches of conspecific eggs or eggs of other congeneric species, i.e. E. ferrugata. The females of E. grisea did not adopt larvae of Elasmostethus interstinctus (Fig. 19); they left after only very short contact with them. In only one case, two E. grisea females with egg batches were found on one leaf in natural conditions. After transfer to the container, they did not react to one another until the larvae hatched. They then showed defensive behaviour of medium intensity, turning to face one another. The more vigorously the larvae moved, the stronger the females’ reaction, with bodies jerking and wings fanning. In the end, one of the females left her larvae and the second female adopted them (Figs 20–22).

Discussion and conclusions Copulation and oviposition. The pattern of copulation observed in the acanthosomatids studied is typical of most of the Pentatomoidea. JORDAN (1958) and FISCHER (2006) described copulation options in Acanthosomatidae in some detail. A female carrying eggs may well be unwilling to copulate. The females of all acanthosomatid species chose oviposition sites as near as possible to fruit of the host plant, facilitating the larvae’s finding food. Much of the literature indicates that E. grisea egg batches are almost exclusively deposited on the lower surface of leaves (JORDAN 1958, MELBER & SCHMIDT 1975a, MAPPES 1994, MAPPES & KAITALA 1994), but our observations found only 57 % (40 out of 70) of batches deposited on the lower surface. ROTH et al. (2006) also mentioned oviposition on the upper side of leaves in E. grisea. Birch leaves move easily in the breeze, switching upper and lower sides and thus exposure to the sun, rain, and predators, a fact that goes some way towards

442 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Behaviour of Acanthosomatidae during oviposition and parental care explaining oviposition on either leaf surface. An identical situation was also found in E. interstinctus. Elasmucha ferrugata deposited all its egg batches on the lower sides of bilberry leaves, which do not change position at all. KOBAYASHI (1953) also described oviposition by this species on the lower side of host plant leaves. However, females of Cyphostethus tristriatus deposited the eggs in all possible positions on Juniperus communis. FISCHER (2006) described the needles of the host plant as the most obvious place for a C. tristriatus clutch. The literature generally appears to indicate that females that care for the eggs deposit fewer of them than females that take no care of their offspring (e.g. TALLAMY & SCHAEFER 1997). In species that stand over their brood, the size of the batch is limited by the body size of the female covering it (MAPPES 1994). Eggs in the centre of the batch are larger than those at the margins in E. ferrugata (MAPPES et al. 1997). In our specimens, the number of eggs in E. ferrugata batches ranged from 33 to 39 (usually 34, 35, and 36, rarely 38 and 39). However, STRAWIÑSKI (1951) gave a broader range for this species, at 17–52, and KOBAYASHI (1953), with the females on Hydrangea paniculata (Sieb.) (Hydrangeaceae), a narrower one, at 20–25. The differences in numbers may depend on the size of the eggs. Similarly, in E. grisea, from 35 to 45 eggs were laid, most often 38, 39, and 40, rarely 35, 36, 42, and 45, while ROTH et al. (2006) counted from 40 to 50 eggs per batch. The body sizes of both Elasmucha species are comparable; the size of the egg batch is therefore similar. In general, the species that do not indulge in parental care deposit more eggs. In Acanthosomatidae, Elasmostethus humeralis Jakovlev, 1883 lays clutches of more than 200 eggs (KUDO 2001) and Elasmostethus atricornis (Van Duzee, 1904) deposits several batches of around 25 eggs each (CARTER & HOEBEKE 2003). The number of batches laid by species with no parental care varies widely. Elasmostethus interstinctus and C. tristriatus deposited a large number of batches with a lower number of eggs, possibly a strategy against egg predators, which either cannot find all the small batches or are replete with several batches and thus leave the remainder untouched. Moreover, Cyphostethus tristriatus has only the limited surface of narrow needles or fruit of Juniperus for its egg batch, and is forced by sheer physical area to divide the eggs into small groups. The highest number of eggs in one batch for C. tristriatus is 14, according to FISCHER (2006) because the females have paired ovaria with seven ovarioles each. However, we also found batches with 20 and 19 eggs. The eggs of both species that leave the eggs without care show a further adaptation to unassisted survival; shortly after deposition, the colour of the eggs turns green, merging with the colour of the background. The eggs of the species in which females care for the eggs are white; they do not change colour, which contrasts with the background. However, these eggs are not visible in the natural “caring” position with the female over the clutch. Acanthosoma haemorrhoidale is an exception; its eggs are whitish even though the species is without maternal care. However, greenish pigmentation is hardly necessary as the batches are always located on the ventral side of a rowan leaf, which is significantly lighter that the dorsal side. The first instar in A. haemorrhoidale and

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Cyphostethus tristriatus is inconspicuously greenish, whereas in Elasmosthetus interstinctus it is red, perhaps as a warning coloration. It has been generally assumed that a female with care invests her time and energy in only one clutch and the care of it (TALLAMY & SCHAEFER 1997). However, more egg batches have also been mentioned in several species, e.g. Sehirus cinctus (Palisot, 1811) from Cydnidae (KIGHT 1998). A second, or even a third, batch can compensate for a decayed batch or provide an increased number of offspring to make the most of optimal conditions (STRAWIÑSKI 1951). KAITALA & MAPPES (1997) recognized that E. ferrugata regularly deposits a second egg batch (the only European Elasmucha species to do so). Our results confirmed this pattern. Elasmucha grisea laid only one batch in central European conditions (also pointed out by FISCHER (2006)). No clear explanation exists as to why oviposition behaviour varies between these species. KAITALA & MAPPES (1997) suggested that parental care lasts a shorter time when eggs are larger and in lower numbers, as the female can deposit further batches. Elasmucha ferrugata admittedly shows a lower number of eggs in its batches than does E. grisea, but only insignificantly so. Parental care. Eight members of the family Acanthosomatidae occur in central Europe. An absence of the parental care has been confirmed in three of them. Elasmostethus interstinctus. Although TEYROVSKÝ (1920) maintained that parental care had not been recognized in central European populations of E. interstinctus, STEHLÍK (1984) mentioned parental care. Other authors who studied the biology of this species found no parental care either (e.g., BUTLER 1923, MACGILL 1942, MAPPES et al. 1996, FISCHER 2006). Our study also recorded an absence of parental care in E. interstinctus. All the egg batches were abandoned by the females, both in the wild and under laboratory conditions. Egg cannibalism was observed. The eggs are green, almost indistinguishable from the background to the human eye, almost certainly representing a defensive adaptation. The absence of parental care is probably a feature of the genus Elasmostethus, as it is not mentioned in other species (e.g. JONES & MCPHERSON 1980; KUDO 1990, 2001; CARTER & HOEBEKE 2003). Cyphostethus tristriatus. BUTLER (1923) and STEHLÍK (1984) recorded that C. tristriatus lacks parental care. FISCHER (2006) described the reproductive behaviour of C. tristriatus in detail for the first time. Like the previous species, C. tristriatus has developed its own strategy. Females deposit larger numbers of egg batches consisting of a lower numbers of eggs. Moreover, females are limited by the space available on the narrow needles and small berries of the host plant , Juniperus communis. Cyphostethus tristriatus also shows adaptation in egg coloration; they are pale green. Acanthosoma haemorrhoidale does not exhibit parental care, as briefly noted by FISCHER (2006). Details of its biology are presented in HANEL & HANELOVÁ (2008). Elasmucha species. All three European Elasmucha species exhibit maternal care. More is known about E. grisea, less about E. ferrugata and E. fieberi. The biology of E. fieberi and E. grisea is similar (MELBER & SCHMIDT 1975a, ROTH et al. 2006, HANEL & HANELOVÁ 2011). Maternal care is one of the features of the entire genus, and is mentioned in all species for which the biology is known. However, the duration of care varies between Elasmucha species.

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We confirmed that E. grisea protects its larvae until the third instar, although MELBER & SCHMIDT (1975a) and MAPPES (1994) extend this to the fourth instar as well, while ROTH et al. (2006) limit it to just the first instar. The explanation of this wide range of data about care in E. grisea is not known. Elasmucha ferrugata continues care until the second instar. A shorter period of care, just for the first instar in E. ferrugata, was recorded by SOUTHWOOD & LESTON (1959), KAITALA & MAPPES (1997) and MAPPES et al. (1997). Care of the first instar was significantly more intensive than that for the second instar in E. ferrugata, which may indicate a trend. However, larvae of the second instar still fed close to the female and spent the night with her. Females of E. fieberi left their larvae during the second instar (MELBER & SCHMIDT 1975a). Wide variability exists in the duration of maternal care within Acanthosomatidae. Only the first instar comes under the protection of Anaxandra gigantea (Matsumura, 1913) after HAYASHI (1987) and Elasmucha grisea, after ROTH et al. (2006). Larvae are protected until the second instar by some females of Anaxandra gigantea and of Sastragala esakii Hasegawa, 1959 after HASEGAWA (1967). Elasmucha signoreti (KUDO & NAKAHIRA 1993), Elasmucha ferrugata (present paper) and Elasmucha fieberi (MELBER & SCHMIDT 1975a) also protect larvae until the second instar. Females of Elasmucha putoni Scott, 1874 (HONBO & NAKAMURA 1985, KUDO 1990, TACHIKAWA 1971), and according to the observations in hand, Elasmucha grisea as well, continue care until the third instar, while those of Elasmucha dorsalis (KUDO et al. 1989, KUDO 1990) and some of the females of Elasmucha grisea according to MELBER & SCHMIDT (1975a) carry on until the fourth instar. Some E. dorsalis females are still exhibiting care routines even during the fifth instar (KUDO et al. 1989, KUDO 1990). Female feeding. Reviewing the literature on whether acanthosomatid females feed or not during their care of offspring gives ambiguous results. Many authors have maintained that the females of a number of species probably fast, e.g. Elasmucha signoreti (KUDO & NAKAHIRA 1993), E. fieberi (MELBER & SCHMIDT 1975a), and E. grisea (MELBER & SCHMIDT 1975a). KAITALA & MAPPES (1997) cited DOLLING (1991) that females of E. ferrugata did not leave the egg batch and only sucked from the mesophyllum. However, KUDO et al. (1989) found that some Elasmucha dorsalis females with eggs and first instars probably fed during their care. We established that E. ferrugata and E. grisea females took nourishment in the period during which they were caring for eggs. They left the clutch and sucked close to it, then returned immediately after feeding. The hypothesis that females fast was offered, by BEQUAERT (1935) among others, but this was not based on exact, direct studies. If a fasting female were to perish, her clutch would be left defenceless. It appears that risking a little time and energy to feed represents a far better investment than leaving the clutch entirely vulnerable to predator or parasitoid should the parent succumb to starvation. Discrimination between broods and adoption. Females of Elasmucha species very probably employ chemical sensitivity when they are search for an egg batch. The acceptance threshold appears to lie at species, rather than individual, level. In all cases, a seeking female adopted the first conspecific clutch that she came upon. It may also be

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 445 J. HANELOVÁ & J. VILÍMOVÁ important how long a female has already stayed with her clutch and how long she has been away from it. It is generally suggested that subsocial hemipterans cannot distinguish their eggs from other conspecific eggs. That situation had previously been confirmed in Elasmucha dorsalis, E. grisea, and E. putoni, and now in E. ferrugata (e.g., MELBER & SCHMIDT 1975a, HONBO & NAKAMURA 1985, FAETH 1989). TSUKAMOTO & TOJO (1992) and FILIPPI-TSUKAMOTO et al. (1995) compared the situation in Elasmucha species with the behaviour of Parastrachia japonensis (Scott, 1880) (Parastrachiidae) with reference to maternal care. A P. japonensis female exhibits “evasive guarding behaviour”; she may remove the eggs from a dangerous location and provide the larvae with berries from the host tree. The female may thus distinguish her own larvae by the “smell” of berries brought to the nest with larvae. In contrast, Elasmucha species glue the egg batch tightly to the surface of the host plant. The female thus defends the place with clutch and food, and her defensive behaviour is more aggressive. In natural conditions, a female does not leave the eggs, thus never coming under any pressure to distinguish her own clutch from any other. Female behaviour. The defensive behaviour of E. grisea females with an egg batch was first mentioned by MODEER (1764). The greatest intensity of defence is triggered by a secretion from the nymphal dorso-abdominal scent glands (e.g. MASCHWITZ & GUTMANN 1979, KUDO et al. 1989, KUDO 1990). The results of our experiments categorised the five types of defensive behaviour already described and classified by MELBER & SCHMIDT (1975a) and complemented by MAPPES et al. (1995), depending on the perceived intensity of attack. The sequence in which these types occur is not always identical, and not all must necessarily be performed. We confirm the results of TEYROVSKÝ (1920), that defensive behaviour differs with the size of aggressor and intensity of the threat. The defensive reaction of an E. ferrugata female with larvae is more intensive than that of one with only eggs. It appears that the female has already made a larger investment in larvae that have hatched than in “merely” the egg batch. One of the intensive reactions is the fanning the wings, which has several possible explanations. The female may be conveying a visual impression of increased body size, or she may be trying, quite literally, to blow away both intruder and unwanted males (TALLAMY & WOOD 1986, Tingidae). However, a female may also be releasing a secretion from the dorso- abdominal glands, more quickly distributed and directed against potential danger by wing-fanning (PODOUBSKÝ 1997). This defensive behaviour of the E. ferrugata female is very similar to that of other Elasmucha species. KUDO et al. (1989) recorded the reaction of an E. dorsalis female visually stimulated by shaking forceps in front of it; KUDO (1990) noted that an E. putoni behaved similarly, as did an E. signoreti female, stimulated by shaking a small black ball at it (KUDO & NAKAHIRA 1993). Comparison with other Pentatomoidea with parental care. Data on parental care of offspring within the Pentatomoidea are sparse. In the temperate zone, females of only two families, Acanthosomatidae and Cydnidae, are known to exhibit this behaviour. Maternal care is known from more taxa in the latter family; it runs in two possible patterns. The female of Cydnus aterrimus (Forster, 1771) cares for the egg batch and feeds the larvae

446 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Behaviour of Acanthosomatidae during oviposition and parental care with a special secretion, probably from the rectum (SCHORR 1957). In the second pattern, the female defends the eggs and provides the larvae with seeds for food; this is more widespread within the Cydnidae, e.g. in Adrisa species (summary in FILIPPI et al. 2001), in Sehirus species (e.g., SITES & MCPHERSON 1982, KIGHT 2000), in Tritomegas species (SOUTHWOOD & LESTON 1959, TALLAMY & WOOD 1986), and others. The pattern described in Cydnidae is very similar to that of Parastrachia japonensis, in the Palaearctic region, the female of which deposits the eggs in a hole she digs in the ground. The female guards the eggs passively, and she may translocate them in the event of threat. After the larvae hatch, the female provides them with food in the form of seeds or berries. The similarity between maternal care in Cydnidae and Parastrachiidae is clear. They live under vegetation, on or in the substratum, assumed to be an ancestral type of biotope for a hypothetical ancestor of the Pentatomoidea (SCHAEFER 1972). Paternal/maternal care could in this light be considered a plesiomorphic feature within Pentatomoidea, at the least. The defensive behaviour of pentatomoids should thus have a similar pattern, as may be observed in the very similar reactions of females to threat in both Elasmucha ferrugata, and Parasrachia japonensis. This pattern may represent an ancestral mode of female behaviour within Pentatomoidea.

Acknowledgements It is our great pleasure to dedicate this paper to Pavel Lauterer and Jaroslav Stehlík, two luminaries of Czech hemipterological research. It is also our very sad obligation to dedicate this paper to Vojtìch Jarošík, recently and suddenly deceased, our colleague and an inspiration to the senior author during her studies. We also extend our gratitude to Lubomír Hanel (Nature Conservation Agency of the Czech Republic, Prague) and Petr Šrámek (Brno) for their help on the field trips and with the photographic documentation. We appreciate the help of Petr Kment (National Museum, Prague) in gathering certain papers together. We thank also Petr Kment and Carl W. Schaefer (University of Connecticut, Storrs) for highly valuable comments on the text and Tony Long (Svinošice) for helping to work up the English.

References

ALDRICH J. R. 1988: Chemical ecology of the Heteroptera. Annual Review of Entomology 33: 211–238. BEQUAERT J. 1935: Presocial behavior among the Hemiptera. Bulletin of Brooklyn Entomological Society 30: 177–191. BOSELLI F. B. 1932: Instinti materni del Sehirus sexmaculatus Rbr. (Heteroptera: Cydnidae. Bolletino del Laboratorio di Zoologia Generale e Agraria del R. Instituto Superiore Agrario in Portici 26: 1–8. BUTLER E. A. 1923: Biology of the British Hemiptera – Heteroptera. H. F. & G. Witherby, London, viii + 682 pp. + 7 pls. CARTER M. E. & HOEBEKE E. R. 2003: Biology and seasonal history of Elasmostethus atricornis (Van Duzee) (Hemiptera: Acanthosomatidae), with descriptions of the immature stages and notes on Pendergrast organ. Proceedings of the Entomological Society of Washington 105: 525–534. COBBEN R. H. 1968: Evolutionary trends in Heteroptera. Part I. Eggs, architecture of the shell, gross embryology and eclosion. Centre for Agricultural Publishing and Documentation, Wageningen, 475 pp.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 447 J. HANELOVÁ & J. VILÍMOVÁ

COUTURIER A. 1946: Remarques sur le curieux comportement d’Elasmucha grisea L. (Hemipt. Pentatom.). L‘Entomologiste 2: 189–192. DOLLING W. R. 1991: The Hemiptera. New York, Oxford University Press, x + 274 pp. DRAKE C. J. 1922: Heteroptera in the vicinity of Cranberry Lake. Pp. 54–86. In: OSBORN H. & DRAKE C. J. (eds.): An ecological study of the Hemiptera of the Cranberry Lake Region, New York. N.Y. State Coll. For. Syracuse Univ. Tech. Publ. 16, Vol. 22, 5, 511 pp. DUDA L. 1879: Zvláštní úkaz mateøské péèe u hmyzu. [Interesting case of maternal care in insects.] Vesmír 8: 15–16 (in Czech). FAETH S. H. 1989: Maternal care in a lace bug, Corythucha hewitti (Hemiptera: Tingidae). Psyche 96: 101–110. FAÚNDEZ E. I. & OSORIO G. A. 2010: New data on the biology of Sinopla perpunctatus Signoret, 1864 (Hemiptera: Heteroptera: Acanthosomatidae). Boletín de Biodiversidad de Chile 3: 24–31. FILIPPI L., HIRONAKA M. & NOMAKUCHI S. 2001: A review of the ecological parametres and implications of subsociality in Parastrachia japonensis (Hemiptera: Cydnidae), a semelparous species that specializes on a poor resource. Population Ecology 43: 41–50. FILIPPI L., HIRONAKA M., NOMAKUCHI S. & TOJO S. 2000: Provisioned Parastrachia japonensis (Hemiptera: Cydnidae) nymphs gain access to food and protection from predators. Animal Behaviour 60: 757–763. FILIPPI-TSUKAMOTO L., NOMAKUCHI S., KUKI K. & TOJO S. 1995: Adaptiveness of parental care in Parastrachia japonensis (Hemiptera: Cydnidae). Annals of the Entomological Society of America 88: 374–383. FISCHER C. 2006: The biological context and evolution of Pendergast’s organs of Acanthosomatidae (Heteroptera, Pentatomoidea). Pp. 1041–1054. In: RABITSCH W. (ed.): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia 19: 1–1184. FROST S. W. & HABER V. R. 1944: A case of parental care in Heteroptera. Annals of the Entomological Society of America 37: 161–166. GUILBERT E. 2003: Habitat use and maternal care of Phloea subquadrata (Hemiptera: Phloeidae) in the Brasilian Atlantic forest (Espirito Santo). European Journal of Entomology 100: 61–63. GOGALA M., YONG H. & BRÜHL C. 1998: Maternal care in Pygoplatys bugs (Heteroptera: Tessaratomidae). European Journal of Entomology 95: 311–315. HANEL L. & HANELOVÁ J. 2007: Ploštice èeledi knìzovitých na Podblanicku. (Faunistic note on the family Acanthosomatidae (Heteroptera, Pentatomomorpha, Insecta) within the Region of Podblanicko (Central Bohemia, Czech Republic.) Sborník Vlastivìdných Prací z Podblanicka 44–47: 71–84 (in Czech, English summary). HANEL L. & HANELOVÁ J. 2008: Co víme o biologii knìze velkého. (What we have known about the Hawthorn Shield Bug’s biology.) Živa 56/94: 30–31 + xvi + p. 3 of cover (in Czech, separate English abstract). HANEL L. & HANELOVÁ J. 2011: Naši knìzové rodu Elasmucha a jejich rodièovské chování. (Native bugs of the genus Elasmucha and their parental care.) Živa 59/97: 27–29 + xvi (in Czech, separate English abstract). HANELOVÁ J. 2005: Péèe o potomstvo u ploštic èeledi Acanthosomatidae (Heteroptera). [Parental care of true bugs from the family Acanthosomatidae (Heteroptera).] Unpublished M.Sc. thesis, Charles University in Prague, Department of Zoology, 103 pp. + 39 photos (in Czech). HASEGAWA H. 1967: Life history of the subsocial bug: egg-guarding behavior of Sastragala esakii. Pp. 145–178. In: IWATA K., HURUKAWA H. & YASUMATSU K. (eds.): Subsocial Insects. Tokyo, Nihon Kontyû-ki IV, Kôdansha, 260 pp. (in Japanese). HAWKINS R. D. 1995: Elasmostethus tristriatus and related shieldbugs in the Sheffield area. Sorby Record 31: 66–67. HAYASHI M. 1987: Parental care observed in Anaxandra gigantea (Heteroptera, Acanthosomatidae). Rostria 38: 569–570. HELLINS J. 1870: A fragment of a life-history of Acanthososma grisea. Entomologist’s Monthly Magazine 7: 53–55. HELLINS J. 1872: Note on the habit of Acanthosoma grisea. Entomologist’s Monthly Magazine 9: 13. HELLINS J. 1874: Additional notes on the egg-laying, etc., of Acanthosoma grisea. Entomologist’s Monthly Magazine 11: 42–43. HONBO Y. & NAKAMURA K. 1985: Effectiveness of parental care in the bug Elasmucha putoni Scott (Hemiptera: Acanthosomatidae). Japanese Journal of Applied Entomology and Zoology 29: 223–229. HUSSEY R. F. 1934: Observations on Pachycoris torridus (Scop.), with remarks on parental care in other Hemiptera. Bulletin of the Brooklyn Entomological Society 29: 133–145.

448 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Behaviour of Acanthosomatidae during oviposition and parental care

JONES W. A. & MCPHERSON J. E. 1980: The first report of the occurrence of acanthosomatids in South Carolina. Journal of the Georgia Entomological Society 15: 286–289. JORDAN K. H. C. 1958: Die Biologie von Elasmucha grisea L. Beiträge zur Entomologie 8: 385–397. KAITALA A. & MAPPESS J. 1992: Evolution of parental care in insects. Luonnon Tutkija 96: 158–162. KAITALA A. & MAPPES J. 1997: Parental care and reproductive investment in shield bugs (Acanthosomatidae, Heteroptera). Oikos 80: 3–7. KAITALA A., HARDLING R., KATVALA M., ORDONEZ R. M. & MIETTINEN M. 2001: Is nonparental egg carrying parental care? Behavioral Ecology 12: 367–368. KIGHT S. L. 1996: Cancaveation and maintenance of maternal behavior in a burrower bug (Sehirus cinctus): A comparative perspective. Journal of Comparative Psychology 110: 69–76. KIGHT S. L. 1998: Precocene II modifies maternal responsiveness in the burrower bug, Sehirus cinctus (Heteroptera). Physiological Entomology 23: 38–42. KIGHT S. L. 2000: Altered photocyclic regimes influence the duration of maternal care in a burrower bug (Heteroptera: Cydnidae). Entomological News 111: 67–73. KIRKALDY G. W. 1904: Upon maternal solicitude in Rhynchota and other nonsocial insects. Annual Report of the Smithsonian Institute 1904: 577–585. KOBAYASHI T. 1953: The development stages of six species of the Japanese Pentatomoidea (Hemiptera). Scientific Reports of the Matsuyama Agricultural Colllege 11: 73–78. KUDO S. 1988: Notes on the oviposition site of Elasmucha putoni (Hemiptera, Acanthosomatidae). Kontyû (Tokyo) 56: 687–688. KUDO S. 1990: Brooding behavior in Elasmucha putoni (Heteroptera: Acanthosomatidae), and a possible nymphal alarm substance triggering guarding responses. Applied Entomology and Zoology 25: 431–437. KUDO S. 1996: Ineffective maternal care of subsocial bug against a nymphal parasitoid: a possible consequence of specialization to predators. Ethology 102: 227–235. KUDO S. 2000: The guarding posture of females in the subsocial bug Elasmucha dorsalis (Heteroptera: Acanthosomatidae). European Journal of Entomology 97: 137–139. KUDO S. 2001: Intraclutch egg-size variation in acanthosomatid bug: adaptive allocation of maternal investment? Oikos 92: 208–214. KUDO S. 2006: Within-clutch egg-size variation in a subsocial bug: the positional effect hypothesis. Canadian Journal of Zoology 84: 1540–1544. KUDO S. & NAKAHIRA T. 1993: Brooding behavior in the bug Elasmucha signoreti (Heteroptera: Acanthosomatidae). Psyche 100: 121–126. KUDO S., SATO M. & OHARA M. 1989: Prolonged maternal care in Elasmucha dorsalis (Heteroptera: Acanthosomatidae). Journal of Ethology 7: 75–81. MACGILL E. 1942: Notes on the early stages of three Pentatomidae (Hem.). Entomologist’s Monthly Magazine 78: 200–202. MAPPES J. 1994: Maternal care and reproductive tactics in shield bugs. Biological Research Report from the University of Jyväskylä 40: 1–30. MAPPES J. & KAITALA A. 1994: Experiments with Elasmucha grisea L. (Heteroptera: Acanthosomatidae): does a female parent bug lay as many eggs as she can defend? Behavior Ecology 5: 314–317. MAPPES J., KAITALA A. & ALATALO R. V. 1995: Joint brood guarding in parent bugs – an experiment on defence against predation. Behavioral Ecology and Sociobiology 36: 343–347. MAPPES J., KAITALA T. & RINNE V. 1996: Temporal variation in reproductive allocation in a shield bug Elasmostethus interstinctus. Journal of Zoology 240: 29–35. MAPPES J., MAPPES T. & LAPPALAINEN T. 1997: Unequal maternal investment in offspring quality in relation to predation risk. Evolutionary Ecology 11: 237–243. MASCHWITZ U. & GUTMANN C. 1979: Spur- und Alarmstoffe bei der gefleckten Brutwanze Elasmucha grisea. Insectes Sociaux 26: 101–111. MELBER A., HÖLSCHER L. & SCHMIDT G. H. 1980: Further studies on the social behaviour and its ecological significance in Elasmucha grisea L. (Hem.-Het.: Acanthosomatidae). Zoologische Anzeiger 205: 27–38. MELBER A. & SCHMIDT G. H. 1975a: Sozialverhalten zweier Elasmucha-Arten. Zeitschrift für Tierpsychologie 39: 403–414. MELBER A. & SCHMIDT G. H. 1975b: Ökologische Bedeutung des Sozialverhaltens zweier Elasmucha-Arten (Heteroptera: Insecta). Oecologia (Berlin) 18: 121–128.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 449 J. HANELOVÁ & J. VILÍMOVÁ

MODEER A. 1764: Några måkvärdigheter hos insectet Cimex ovatus pallidegriseus, abdominis lateribus albo nigroque variis albis, basi scutelli nigricante. Kongliga Svenska Vetenskaps Akademiens Handlingar 25: 41–57. [Only cited after TALLAMY (1999)]. MONTEITH G. B. 2006: Maternal care in Australian oncomerine shield bugs (Insecta, Heteroptera, Tessaratomidae). Pp. 1135–1152. In: RABITSCH W. (ed.): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia 19: 1–1184. NOMAKUCHI S., FILIPPI L. & TOJO S. 1998: Selective foraging behavior in nest-provisioning females of Parastrachia japonensis (Hemiptera: Cydnidae): Cues for preferred food. Journal of Insect Behavior 11: 605–619. ODHIAMBO T. R. 1960: Parental care in bugs and non-social insects. New Scientist 18: 449–451. OGORZA£EK A. & TROCHIMCZUK A. 2009: Ovary structure in a presocial insect, Elasmucha grisea (Heteroptera: Acanthosomatidae). Arthropod Structure & Development 38: 509–519. PARR A., TALLAMY D. W., MONACO E. L. & PESEK J. D. 2002: Proximate factors regulating maternal options in the eggplant lace bug, Gargaphia solani (Heteroptera: Tingidae). Journal of Insect Behaviour 15: 495–511. PIERRE 1903: Note sur les murs d’ Elasmostethus griseus Linn. = interstinctus Reut. (Hémipt.). Bulletin de la Société Entomologique de France 1903: 131–132. PODOUBSKÝ M. 1997: Srovnávací morfologická studie dorsálních abdominálních pachových žláz zástupcù nadèeledi Pentatomoidea (Heteroptera). [Comparative morphological study of dorsal abdominal scent glands in Pentatomoidea (Heteroptera).] Unpublished M.Sc. thesis, Charles University in Prague, Department of Zoology, 58 pp. (in Czech). PRUNER L. & MÍKA P. 1996: Seznam obcí a jejich èástí v Èeské republice s èísly mapových polí pro síťové mapování fauny. (List of settlements in the Czech Republic with associated map field codes for faunistic grid mapping system). Klapalekiana 32 (Suppl.): 1–115 (in Czech, English summary). RIDLEY M. 1978: Paternal care. Animal Behaviour 26: 904–932. ROSSUM A. J. 1904: Note on Elasmostethus. Entomologische Berichten (Amsterdam) 21: 204–206. ROTH S., ADASCHKIEWITZ W. & FISCHER C. 2006: Notes on the bionomics of Elasmucha grisea (Linnaeus, 1758) (Heteroptera, Acanthosomatidae) with special regard to joint guarding. Pp. 1153–1167. In: RABITSCH W. (ed.): Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia 19: 1–1184. SCHAEFER C. W. 1972: Degree of metathoracic scent-gland development in the trichophorous Heteroptera (Hemiptera). Annals of the Entomological Society of America 65: 810–821. SCHORR H. 1957: Zur Verhaltensbiologie und Symbiose von Brachypelta aterrima Först. (Cydnidae, Heteroptera). Zeitschrift für Morphologie und Ökologie der Tiere 45: 561–602. SITES R.W. & MCPHERSON J. E. 1982: Life history and laboratory rearing of Sehirus cinctus cinctus (Hemiptera: Cydnidae), with descriptions of immature stages. Annals of the Entomological Society of America 75: 210–215. SMITH R. L. 1997: Evolution of paternal care in the giant water bugs (Heteroptera: Belostomatidae). Pp. 116–149. In: CHOE J. C. & CRESPI B. J. (eds.): The evolution of social behavior in insects and arachnids. Cambridge, Cambridge University Press, 541 pp. SOUTHWOOD T. R. E. 1956: The structure of the eggs of the terrestrial Heteroptera and its relationship to the classification of the group. Transactions of the Royal Entomological Society of London 108: 163–221. SOUTHWOOD T. R. E. & LESTON D. 1959: Land and water bugs of the British Isles. London & New York, Frederick Warne & Co. Ltd., ix + 436 pp. STEHLÍK J. L. 1984: Results of the investigations on Hemiptera in Moravia made by the Moravian Museum. Pentatomoidea III. Acta Musei Moraviae, Scientiae Naturales 69: 163–185. STRAWIÑSKI K. 1951: Introductory studies on the biology of Elasmucha ferrugata F. (Hem. - Heteroptera). Annals of the University of Marie Curie-Sk³odowska Lublin (C) 4: 149–163. TACHIKAWA S. 1971: Studies on parental care of Heteroptera I. On Elasmucha putoni of Japan and other Elasmucha spp. Journal of Agricultural Science, Tokyo Nogyo Daiga 18: 24–34. TALLAMY D. W. 1999: Child care among the insects. Scientific American 280: 72–77. TALLAMY D. W. 2000: Sexual selection and the evolution of exclusive parental care in Arthropods. Animal Behaviour 60: 559–567. TALLAMY D. W. 2001: Evolution of exclusive paternal care in Arthropods. Annual Review of Entomology 46: 139–165.

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TALLAMY D. W. & SCHAEFER C. W. 1997: Maternal care in the Hemiptera: ancestry, alternatives and current adaptive value. Pp. 94–115. In: CHOE J. C. & CRESPI B. J. (eds.): The evolution of social behavior in insects and arachnids. Cambridge University Press, Cambridge, 541 pp. TALLAMY D. W. & WOOD T. K. 1986: Convergence pattern in social insects. Annual Review of Entomology 31: 369–390. TEYROVSKÝ V. 1918: Nové poznatky o plošticích. [New data about true bugs.] Èasopis Èeské Spoleènosti Entomologické 15 (Vìstník): xi–xii (in Czech). TEYROVSKÝ V. 1920: Studie o èeských Acanthosominách I. [A study about Czech Acanthosomatinae I.] Èasopis Èeské Spoleènosti Entomologické 17: 11–15 (in Czech). THOMAS D. B. 1991: The Acanthosomatidae (Heteroptera) of North America. Pan-Pacific Entomologist 67: 159–170. TSUKAMOTO L. & TOJO S. 1992: A report of progressive provisioning in a stink bug, Parastrachia japonensis (Hemiptera: Cydnidae). Journal of Ethology 10: 21–29.

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Table 1. Maternal care in the family Acanthosomatidae. (Continued on pages 453–454.)

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Table 2. Number of studied Acanthosomatidae specimens used in the behavioral experiments.

Experiments in Experiments in Reared from SPECIES containers natural conditions egg to adult

Acanthosoma haemorrhoidale 2 adults 20 larvae 4 specimens (Linnaeus, 1758)

Cyphostethus tristriatus 7 adults 20 larvae 10 specimens (Fabricius, 1758)

Elasmostethus interstinctus 25 adults, 10 larvae 30 adults 15 specimens (Linnaeus, 1758)

Elasmucha ferrugata 50 adults 120 larvae + adults 10 specimens (Fabricius, 1787)

Elasmucha fieberi 2 adults 8 larvae 2 specimens Jakovlev, 1864

Elasmucha grisea 55 adults, 15 larvae 150 larvae + adults 20 specimens (Linnaeus, 1758)

Table 3. Host plants of studied Central European Acanthosomatidae.

Bug species Host plant species

Acanthosoma haemorrhoidale Sorbus aucuparia L. (Linnaeus, 1758)

Cyphostethus tristriatus Juniperus communis L. (Fabricius, 1758)

Elasmostethus interstinctus Betula pendula Roth (Linnaeus, 1758) Alnus glutinosa (L.) Gaertn.

Elasmucha ferrugata Vaccinium myrtilus L. (Fabricius, 1787) Lonicera xylosteum L.

Elasmucha fieberi Betula pendula Roth Jakovlev, 1864

Elasmucha grisea Betula pendula Roth (Linnaeus, 1758) Alnus glutinosa (L.) Gaertn. Crataegus sp.

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Table 4. Number of eggs in the individual batches of Acanthosomatidae.

SPECIES Number of eggs Number of batches

Acanthosoma haemorrhoidale 23 1 (Linnaeus, 1758) (N = 6) 28 1 14 1 27 1 16 1 21

Cyphostethus tristriatus 20 1 (Fabricius, 1758) (N = 4) 19 1 31 71

Elasmostethus interstinctus 41 (Linnaeus, 1758) (N = 16) 5 1 61 72 91 11 1 12 2 13 1 18 1 20 1 26 1 27 1 29 1 30 1

Elasmucha ferrugata 33 3 (Fabricius, 1787) (N = 25) 34 5 35 6 36 5 37 3 38 2 39 1

Elasmucha fieberi 34 1 Jakovlev, 1864

Elasmucha grisea 35 1 (Linnaeus, 1758) (N = 20) 36 1 37 2 38 3 39 4 40 3 41 2 42 1 43 2 45 1

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Table 5. Reaction of Elasmucha ferrugata (Fabricius, 1787) female to the attack by soft tweezer. A = antennae straightening; B = antennae jerking; C = body tilting; D = body jerking; E = body swinging; F = wings fanning. 0 = no reaction of female on the convoluted leaf. N = 100 attacks of 20 females.

Combination of reactions Number of answers

A2

A+ B 2

A + B + C 17

A + B + C + D 20

A + B + C + D + E 5

A + B + C + D + E + F 6

A + C + D 37

A + C + D + E 3

A + C + D + F 2

A + C + F 1

A + C + D + F + D 4

01

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 459–493, 2013

New data on the Heteroptera fauna of La Palma, Canary Islands (Insecta: Hemiptera)

BEREND AUKEMA1, HANS DUFFELS2, HANNES GÜNTHER3, CHRISTIAN RIEGER4 & GERHARD STRAUß 5 1Naturalis Biodiversity Center, P.O. Box 9617, NL-2300 RA Leiden, the Netherlands; e-mail: [email protected] 2 Naturalis Biodiversity Center, P.O. Box 9617, NL-2300 RA Leiden, the Netherlands; e-mail: [email protected] 3 Eisenacher Straße 25, D-55218 Ingelheim, Germany; e-mail: [email protected]

4 Lenbachstraße 11, D-72622 Nürtingen, Germany; e-mail: [email protected]

5 Mozartstraße 4, D-88400 Biberach, Germany; e-mail: [email protected]

AUKEMA B., DUFFELS H., GÜNTHER H., RIEGER CH. & STRAUß G. 2013: New data on the Heteroptera fauna of La Palma, Canary Islands (Insecta: Hemiptera). In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 459–493. – Heteroptera collected on La Palma in the Canary Islands during the period 2001–2012 are listed. Data for 153 species are given, including 28 species recorded for the first time from La Palma. Six of these species are also new for the Canary Islands: Cantacader quadricornis (Lepeletier et Serville, 1828) (Tingidae), Bryocoris pteridis (Fallén, 1807), Macrolophus pygmaeus (Rambur, 1839), Thiomiris sulphureus (Reuter, 1879) (all Miridae), Amphiareus obscuriceps (Poppius, 1909) (Anthocoridae) and Ischnodemus quadratus Fieber, 1837 (Lygaeidae). Furthermore, the occurrence of eight species on La Palma is confirmed. With this contribution the number of species known from the Canary Islands and La Palma increases to 400 and 216 respectively. The number of Canarian endemic species on La Palma is 48 (22.2 %). Lindbergopsallus lauri Rieger, 2012 and L. lunariae Rieger, Strauß et Günther, 2011 (Miridae), Collartida tanausu J. Ribes, Oromi et E. Ribes, 1998 (Reduviidae) and Nysius gloriae Baena et Garcia, 1999 (Lygaeidae) are only known from La Palma.

Keywords. Heteroptera, true bugs, distribution, new records, check-list, Palaearctic Region, Canary Islands

Introduction Systematic study of the Heteroptera fauna of the Canary Islands began with the publications of Maurice NOUALHIER (1893) and Håkan LINDBERG (1953, 1960). Thereafter, many heteropterists have collected on the Canary Islands and reported numerous species. Based on published information, AUKEMA et al. (2006) collated the occurrence of species on the different islands. Doubtful and mistakenly listed species were omitted and new records listed, together with collecting data. They reached a total of 385 species for the archipelago and 186 species for La Palma. Since then, five new species have been described: The tingid Kalama montisclari from Montaña Clara Island by J. RIBES & PAGOLA-CARTE (2008) and the mirids Systellonotus stysi J. Ribes, Pagola- Carte et Heiss, 2008 and Atomoscelis pictifrons J. Ribes, Pagola-Carte et Heiss, 2008

459 B. AUKEMA ET AL. from Tenerife by J. RIBES et al. (2008), and Lindbergopsallus lunariae Rieger, Strauß et Günther, 2011 by RIEGER et al. (2011) and L. lauri by RIEGER (2012) from La Palma. Furthermore, Peirates stridulus (Fabricius, 1787), Pasira basiptera Stål, 1859 and Pygolampis bidentata (Goeze, 1778) were listed from the Canary Islands by PUTSHKOV & MOULET (2010) and AUKEMA et al. (2013) added Orsillus depressus (Mulsant et Rey, 1852). Additional island records were listed by AUKEMA (2012), Coranus kiritshenkoi Bergevin, 1932 from Fuerteventura, FARACI (2011), Sigara (Halicorixa) selecta (Fieber, 1848) from Fuerteventura, and SANTAMARÍA et al. (2012), Heliocorisa vermiculata (Puton, 1874) from Tenerife, Mesovelia vittigera Horváth, 1895 from El Hierro and La Gomera, Merragata hebroides F. B. White, 1877 from La Gomera, and Microvelia (Microvelia) gracillima Reuter, 1882 from Fuerteventura and Lanzarote. Two changes were made in the 2006 checklist (see also Table 2): Cantacader divisus Bergroth, 1908 was listed by B. LIS (2003) instead of C. quadricornis (Lepeletier et Serville, 1828) and the record of Liolobus pallidicornis (Reuter, 1891) in HEISS (1997) was based on misidentified Anepsciocoris encaustus (Puton, 1869) (E. HEISS, pers. comm.). To date, 395 species are reported from the Canary Islands. With 294 species, Tenerife is the most species-rich island of the archipelago, followed by Gran Canaria with 235 species. From La Palma, 189 species have been reported to date. We dedicate the paper to our friends and highly esteemed colleagues Jaroslav L. Stehlík, on his 90th birthday, and Pavel Lauterer for his 80th birthday.

Material and methods In this contribution the results of field trips by the authors in the years 2001, 2005, 2007, 2008, 2009, 2011 and 2012, together with some additional records, are provided. Details of the collecting sites are listed in Table 1. The following abbreviations are used for collectors and collections: BA – B. Aukema; CR – Chr. Rieger; GS – G. Strauss; HG – H. Günther; JD – J. P. Duffels; ZMAN – Zoölogisch Museum, Amsterdam (now housed in Naturalis Biodiversity Center, Leiden). Unless stated otherwise (between brackets) the material is deposited in the collection of the collector(s).

New records for the Canary Islands from La Palma

TINGIDAE Cantacader quadricornis (Lepeletier et Serville, 1828) Lomo Oscuro 15.iii.2005, 3 ♂♂ 3 ♀♀, HG; 9.v.2009, 4 ♂♂ 4 ♀♀ 2 larvae 5→2 ♀♀, HG, CR, GS; 7.vi.2011, 1 ♂, GS; 11.vi.2011, 1 ♂, GS.

MIRIDAE Bryocoris pteridis (Fallén, 1807) Fuencaliente, 2.v.2009, 1 ♂ on fern, HG.

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Macrolophus pygmaeus (Rambur, 1839) Los Llanos, 8.v.2009, 1 ♀, at light, CR.

Thiomiris sulphureus (Reuter, 1879) Volcán Teneguía, 30.iv.2009, 1 ♂, on roadside vegetation, HG.

ANTHOCORIDAE Amphiareus obscuriceps (Poppius, 1909) La Lomadita, 21.iii.2010, 2 ♀♀, CR.

LYGAEIDAE Ischnodemus quadratus Fieber, 1837 Bco. de las Angustias, 16.iv.2012, 1 ♂ 1 ♀, on roadside vegetation, HG.

New records for La Palma

TINGIDAE Tingis (Tropidocheila) ajugarum (Frey-Gessner, 1872) La Punta, 24.iii.2009, 6 ♂♂ 3 ♀♀ with larvae; 29.iii.2009, 1 ♂ 2 ♀♀; 2.iv.2009, 6 ♂♂ 5 ♀♀ with larvae, on Ajuga chamaepitys, BA.

MIRIDAE Nesidiocoris tenuis (Reuter, 1895) Faro de Barlovento, 21.iii.2010, 5 ♂♂ 6 ♀♀, on Hyoscyamus niger, CR.

Pinalitus parvulus (Reuter, 1897) Bco. de las Angustias, 5.v.2009, 1 ♂, CR. Fuencaliente NE, 4.v.2009, 2 ♂♂, CR.

Canariocoris hyperici Lindberg, 1951 Bco. de las Angustias, 5.v.2009, 1 ♂ 2 ♀♀, GS; 3.vi.2011, 1 ♂ 1 ♀, GS. Bco. del Jorado 13.vi.2011, 3 ♂♂ 12 ♀♀, GS. Los Barros, 31.iii.2009, 1 ♀, BA.

Canariocoris punctatus (Noualhier, 1893) Playa de Nogales, 7.iii.2008, 1 ♂, JD (ZMAN). Puntagorda, 10.iii.2008, 1 ♂ 2 ♀♀, JD (ZMAN).

Orthotylus (Melanotrichus) flavosparsus (C. R. Sahlberg, 1841) Garafia, 17.iii.2008, 10 ♂♂ 8 ♀♀, JD (ZMAN). La Lomadita, 21.iii.2010, 1 ♂ 3 ♀♀, CR, GS. La Punta, 2.iv.2009, 1 ♀, BA. Puerto Naos, 24.iii.2010, 6 ♂♂ 12 ♀♀, CR, GS; 29.iii.2010, 1 ♂ 1 ♀, CR, GS. Puntagorda, Puerto, 25.iii.2010, 2 ♂♂, CR. Tazacorte, 21.iii.2010, 1 ♀, GS.

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Compsidolon (Coniortodes) cytisi (Lindberg, 1953) Bco de las Angustias, 5.v.2009, 4 ♀♀, CR.

Lindbergopsallus laureti (Lindberg, 1936) Bco. de la Galga, 1.v.2009, 17 ♂♂ 15 ♀♀, HG, CR, GS; 28.iii.2010, 1 ♂ 3 ♀♀, on Laurus, GS. La Lomada, 18.vi.2007, 1 ♀, HG. Laguna Barlovento, 1.iv.2009, 4 ♂♂ 4 ♀♀, BA. Montaña de la Breña, 28.iii.2009, 4 ♂♂ 6 ♀♀, on Myrica faya, BA. Pico Birigoyo, 9.vi.2011, 7 ♂♂ 2 ♀♀, GS.

ANTHOCORIDAE Anthocoris nemoralis (Fabricius, 1794) Puerto de Tazacorte, 13.iii.2008, 1 ♀, JD (ZMAN). Puerto Naos, 24.iii.2010, 4 ♂♂ 8 ♀♀ 1 larva 5, CR, GS; 26.iii.2010, 5 ♂♂ 8 ♀♀ 1 larva 5, on Ricinus, CR.

Orius (Dimorphella) albidipennis (Reuter, 1884) Puerto de Tazacorte, 13.iii.2008, 1 ♂ 1 ♀, JD (ZMAN).

Orius (Orius) niger (Wolff, 1811) Punta Baja de San Simón, 2.iii.2010, 1 ♂, CR.

Cardiastethus nazarenus Reuter, 1884 Montaña de la Breña, 28.iii.2009, 1 ♂ 1 ♀, BA.

Lyctocoris (Lyctocoris) campestris (Fabricius, 1794) Fuencaliente, 14.ii.2001, 1 ♂, HG.

LYGAEIDAE Geocoris (Geocoris) phaeopterus (Germar, 1838) Bco. de la Galga, 12.vi.2007, 1 ♀, HG. Bco. Hondo, 2.v.2009, 2 ♀♀, HG.

Neocamptotelus aeonii (Lindberg, 1953) Bco. Hondo, 30.iv.2009, 4 ♂♂ 1 ♀, HG; 2.v.2009, 28 ♂♂ 18 ♀♀, HG, CR, GS. Fuencaliente N, 7.v.2009, 9 ♂♂ 8 ♀♀, CR, GS; 10.v.2009, 1 ♂ 1 ♀, CR, under Polycarpaea. Lomo Oscuro, 9.v.2009, 1 ♀, CR. Montaña Azufre, 27.iii.2010, 5 ♂♂ 2 ♀♀, CR. Montaña la Laguna, 12.vi.2007, 1 ♂ 1 ♀, HG. Salinas, 12.vi.2007, 1 ♂ 1 ♀, HG.

BERYTIDAE Gampsocoris punctipes punctipes (Germar, 1822) Bco. de las Angustias, 16.iv.2012, 1 ♂ 1 ♀, on Ononis, HG. Nogales, 6–7.iii.2008, 3 ♂♂ 1 ♀, JD (ZMAN). Santo Domingo, 21.iii.2010, 1 ♂, GS.

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COREIDAE Haploprocta incognita Heiss et Moulet, 1994 (Figs 2–3) Torre del Time, 1.vi.2011, 1 ♂ 2 ♀♀, GS.

Gonocerus insidiator (Fabricius, 1787) Tijarafe, Cemeterio, 23.iii.2009, 1 ♀, BA.

Plinachtus imitator (Reuter, 1891) Bco. de las Angustias, 3.vi.2011, 7 ♂♂ 10 ♀♀, GS.

CYDNIDAE Geotomus punctulatus (A. Costa, 1847) Bco. de la Galga, 10.iii.2005, 2 ♂♂ 2 ♀♀, HG; 14.iii.2005, 4 ♂♂ 1 ♀, HG; 12.vi.2007 1 ♂ 2 ♀♀, HG; 18.vi.2007, 4 ♀♀, HG; 1.v.2009, 2 ♂♂ 2 ♀♀, HG, CR. El Paso, farmland, 7.v.2009, 2 ♂♂ 3 ♀♀, 22.iii.2010, 2 ♂♂, 22.iii.2010 2 ♂♂, GS. El Pilar, 3.v.2009, 2 ♂♂ 5 ♀♀, HG, CR; 23.iii.2010, 1 ♀, CR; Laguna de Barlovento, 1.iv.2009, 2 ♂♂ 1 ♀, BA, 21.iv.2012, 1 ♂ 1 ♀, HG. La Lomada, 18.vi.2007, 4 ♀♀. Lomo Oscuro, 9.vi.2007, 5 ♂♂ 5 ♀♀. Montaña de la Breña, 28.iii.2009, 6 ♂♂ 5 ♀♀, BA. Pico de la Nieve, 6.v.2009, 1 ♂ 1 ♀, HG; 24.iv.2012, 1 ♀, HG. San Isidro, 6.v.2009, 1 ♀, CR. Villa de Mazo, 28.iii.2009, 4 ♂♂ 3 ♀♀, BA.

PENTATOMIDAE Acrosternum heegeri Fieber, 1861 Lomo Oscuro, 9.v.2009, 2 ♀♀, GS.

Acrosternum millierei (Mulsant et Rey, 1866) Puerto Naos, 29.iii.2010, 1 ♂, CR.

Confirmed records from La Palma

HEBRIDAE Merragata hebroides F. B. White, 1877 Los Llanos de Aridane, water basin, 20.iii.2009, 4 ♂♂ 3 ♀♀, BA.

TINGIDAE Tingis (Tropidocheila) insularis (Horváth, 1902) Torre del Time, 1.vi.2011, 5 ♂♂ 2 ♀♀, GS.

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NABIDAE Nabis (Nabis) punctatus punctatus A. Costa, 1847 Lomo Oscuro, 20.iii.2010, 1 ♂, GS.

ANTHOCORIDAE Elatophilus pilosicornis Lindberg, 1953 Fuencaliente N, 4.v.2009, 1 ♀, CR. Las Tricias, 8.v.2009, 1 ♀, CR.

Ectomocoris (Ectomocoris) chiragra (Fabricius, 1803) Nogales, 6.–7.iii.2008, 1 ♂, JD (ZMAN).

REDUVIIDAE Sastrapada baerensprungi (Stål, 1859) Bco. de la Galga, 24.ii.2001, 2 ♂♂, HG; 14.iii.2005, 1 ♀, HG. Laguna de Barlovento, 21.iv.2012, 1 ♂ 1 ♀ 1 larva, HG.

COREIDAE Gonocerus juniperi Herrich-Schaeffer, 1839 Tijarafe, Cemeterio, 23.iii.2009, 2 ♀♀, on Cupressus, BA.

ACANTHOSOMATIDAE Cyphostethus tristriatus (Fabricius, 1787) Tijarafe, Cemeterio, 23.iii.2009, 1 ♀, on Cupressus, BA.

Additional records of species already known from La Palma

CORIXIDAE Corixa affinis Leach, 1817 El Paso, gravel pit, 5.vi.2011, 5 ♂♂ 6 ♀♀, GS. Los Llanos de Aridane, water basin, 20.iii.2009, 1 ♀, BA.

Sigara (Vermicorixa) lateralis (Leach, 1817) El Paso, gravel pit, 25.iii.2010, 4 ♂♂ 3 ♀♀, CR; 27.iii.2010, 1 ♂, CR. Los Llanos de Aridane, water basin, 20.iii.2009, 1 ♂ 11 ♀♀, BA.

NOTONECTIDAE Anisops debilis canariensis Noualhier, 1893 El Paso, gravel pit, 25.iii.2010, 2 ♂♂ 2 ♀♀, CR; 27.iii.2010, 2 ♂♂ 5 ♀♀, CR; 6.v.2011, 13 ♂♂ 12 ♀♀, GS. Los Llanos de Aridane, water basin, 20.iii.2009, 3 ♀♀, BA.

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Anisops sardeus sardeus Herrich-Schaeffer, 1849 El Paso, gravel pit, 25.iii.2010, 1 ♀, CR. Los Llanos de Aridane, water basin, 20.iii.2009, 1 ♂ 1 ♀, BA.

TINGIDAE Dictyla indigena (Wollaston, 1858) Bco. del Jorado, 13.vi.2011, 1 ♂ 3 ♀♀, GS. Bco. Herradura, 12.vi.2011, 1 ♂ 2 ♀♀, GS. Bco. Hondo, 30.iv.2009, 2 ♂♂ 2 ♀♀ 1 larva, CR; 2.v.2009, 1 ♀, CR. El Paso, gravel pit, 22.iii.2010, 1 ♂, GS. Fuencaliente N, 7.v.2009, 8 ♂♂ 4 ♀♀, CR, GS; 10.v.2009, 5 ♂♂ 4 ♀♀, CR, GS; 20.iii.2010, 1 ♂, CR, GS. Jedey, 11.vi.2011, 2 ♂♂ 2 ♀♀, GS. La Fajana, 9.iii.2008, 2 ♀♀, JD (ZMAN). La Punta, 2.iv.2009, 4 ♂♂ 5 ♀♀, BA. Las Tricias, 8.v.2009, 4 ♂♂, CR, GS. Lomo Oscuro, 15.iii.2005, 2 ♂♂ 2 ♀♀, HG; 9.v.2009, 4 ♂♂ 2 ♀♀, HG; 20.iii.2010, 2 ♂♂ 2 ♀♀, CR; 7.vi.2011, 1 ♂, GS. Montaña Azufre, 27.iii.2010, 1 ♂ 1 ♀, CR. Montaña la Laguna, 2.v.2009, 6 ♂♂ 4 ♀♀, CR, GS. Monte de Luna,18.iv.2000, 1 ♂ 1 ♀, H. Ziegler leg. (CR). Nogales, 6–7.iii.2008, 1 ♂ 3 ♀♀, JD (ZMAN). Punta Baja de San Simón, 4.v.2009, 3 ♂♂ 13 ♀♀, CR, GS. Puntagorda, Puerto, 25.iii.2010, 8 ♂♂ 2 ♀♀ 4 larvae, CR. San Simón, 27.iii.2010, 1 ♀, CR. Tigalate, 14.iv.2000, 1 ♀, H. Ziegler leg. (CR). Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS. Volcán Martín, 10.v.2009, 1 ♂, HG. Volcán Teneguía, 30.iv.2009, 1 ♂ 2 ♀♀, HG, CR.

Dictyla nassata (Puton, 1874) Bco. del Jorado, 23.iii.2009, 1 ♂, BA; 13.vi.2011, 1 ♂, GS. Bco. Herradura, 12.vi.2011, 1 ♂, GS. Bco. Hondo, 30.iv.2009, 4 ♂♂ 4 ♀♀, CR, GS; 2.v.2009, 1 ♀, HG. El Paso, farmland, 7.v.2009, 2 ♂♂ 2 ♀♀, HG, CR, GS; 11.v.2009, 1 ♂ 1 ♀, HG, GS. El Pinar, 25.iii.2009, 1 ♂, BA. Fuencaliente N, 7.v.2009, 2 ♂♂ 2 ♀♀, CR, GS. Garafia, 17.iii.2008, 1 ♀, JD (ZMAN). Las Tricias, 8.v.2009, 2 ♀♀, GS. Montaña de la Negra, 25.iii.2010, 1 ♂, CR. Nogales, 6.–7.iii.2008, 1 ♂ 1 ♀, JD (ZMAN). Puntagorda, 17.iii.2008, 2 ♂♂ 1 ♀, JD (ZMAN); 7.iv.2000, 1 ♀, H. Ziegler leg. (CR). Tigalate, 14.iv.2000, 1 ♀, H. Ziegler leg. (CR). Volcán Martín, 10.v.2009 1 ♀, HG. Volcán San Antonio, 30.iii.2009, 1 ♂, BA.

Tingis (Tingis) canariensis Péricart, 1981 Bco. de los Angustias, 30.iii.2009, 1 ♀, 31.iii.3009, 1 ♂ 14 ♀♀, on Carlina salicifolia, BA. San Nicolás, 20.iii.2010, 30 ♂♂ 50 ♀♀, CR, GS; 24.iii.2010, 22 ♂♂ 18 ♀♀, on Carlina salicifolia, CR, GS.

Tingis (Tingis) denudata Horváth, 1906 Bco. Hondo, 2.v.2009, 1 ♂, HG. La Punta, 2.iv.2009, 5 ♂♂ 1 ♀, on Centaurea melitensis, BA.

Tingis (Tingis) maderensis (Reuter, 1890) Bco. Herradura, 12.vi.2011, 1 ♂, GS. El Palmar, 27.iii.2009, 1 ♀, BA.

MIRIDAE Cyrtopeltis canariensis (Lindberg, 1936) Barlovento, 21.iii.2010, 3 ♂♂, CR, GS. Bco. del Jorado, 23.iii.2009, 11 ♂♂ 7 ♀♀, BA; 13.vi.2011, 2 ♂♂ 1 ♀, GS. Bco. de las Angustias, 5.v.2009, 1 ♂ 1 ♀, GS. El Paso, farmland, 11.vi.2007, 1 ♀, HG; 7.v.2009, 5 ♂♂ 2 ♀♀, CR, GS; 11.v.2009, 6 ♂♂, HG; 22.iii.2010, 1 ♂ 1 ♀, on Cistus symphytifolius, CR. El Pilar, 17.vi.2007, 11 ♂♂ 2 ♀♀, HG; 21.iii.2009, 1 ♂, BA; 3.v.2009, 3 ♂♂ 2 ♀♀, HG, GS. Hoya Grande, 8.vi.2011, 1 ♀, GS. Las Tricias, 6.v.2009, 1 ♂, GS. Pico de la Nieve, 1070 m a.s.l., 6.v.2009, 13 ♂♂ 10 ♀♀, CR, GS. Puerto Naos, 24.iii.2010, 1 ♀, GS. Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 465 B. AUKEMA ET AL.

Dicyphus (Dicyphus) rubicundus Blöte, 1929 Bco. de las Angustias, 5.v.2009, 25 ♂♂ 7 ♀♀, on Grenovia aurea, HG, CR, GS. Las Tricias, 7.v.2009, 3 ♂♂ 3 ♀♀, CR.

Macrolophus melanotoma (A. Costa, 1853) Bco. Herradura, 12.vi.2011, 1 ♂ 1 ♀, GS. Las Tricias, 7.v.2011, 3 ♂♂ 2 ♀♀, GS.

Macrolophus pericarti J. Ribes et Heiss, 1998 El Pilar, 21.iii.2009, 12 ♀♀, on Cistus symphytifolius, BA.

Deraeocoris (Camptobrochis) serenus (Douglas et Scott, 1868) Bco. del Jorado, 10.vi.2011, 1 ♀, GS. Bco. de las Angustias, 6.vi.2007, 1 ♀. Bco. Hondo, 30.iv.2009, 3 ♀♀ 1 larva, CR, GS; 2.v.2009, 2 ♀♀, CR. El Pinar, 25.iii.2009, 1 ♂, BA. Fuencaliente NE, 4.v.2009, 1 ♀, CR. Garafia, 17.iii.2008, 4 ♂♂ 4 ♀♀, JD (ZMAN). Jedey, 11.vi.2011, 1 ♀, GS. La Lomadita, 21.iii.2010, 2 ♂♂ 2 ♀♀, CR, GS. Las Tricias, 7.v.2011, 1 ♀, GS. Lomo Oscuro, 7.vi.2011, 1 ♂ 1 ♀, GS. Los Llanos, 6.vi.2007, 1 ♀, HG. Montaña de la Negra, 25.iii.2010, 1 ♂, CR. Puerta de Tazacorte, 14.iii.2008, 1 ♂ 1 ♀, JD (ZMAN). Puerto Naos, 24.iii.2010, 1 ♂ 1 ♀, CR, GS. Puntagorda, 10.iii.2008, 2 ♀♀, JD (ZMAN). Roque de los Muchachos, 27.ii.2001, 2 ♂♂, HG. Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS.

Eurystylus bellevoyei (Reuter, 1879) La Lomadita, 21.iii.2010, 4 ♂♂ 19 ♀♀, CR. Tazacorte, 21.iii.2010, 1 ♀, GS.

Pinalitus insularis (Reuter, 1895) Bco. de la Galga, 24.ii.2001, 1 ♀, HG; 1.v.2009, 5 ♂♂ 9 ♀♀, on Laurus, HG, CR, GS. Bco. de las Angustias, 5.v.2009, 1 ♀, HG; 3.vi.2011, 2 ♂♂ 1 ♀, GS. Bco. Herradura, 12.vi.2011, 1 ♂, GS. El Pilar, 21.iii.2009, 1 ♂, BA; 3.v.2009, 1 ♀, HG. Fuencaliente NE, 4.v.2009, 3 ♀♀, CR, GS. Garafia, 17.iii.2008, 1 ♀, JD (ZMAN). La Lomada, 18.vi.2007, 1 ♂ 10 ♀♀, HG. La Lomadita, 21.iii.2010, 1 ♂, GS. Laguna de Barlovento, 1.iv.2009, 1 ♂ 4 ♀♀, on Laurus, BA. Las Tricias, 8.v.2009, 1 ♂ 2 ♀♀, CR. Lomo Oscuro, 9.vi.2007, 1 ♂ 2 ♀♀, HG; 9.v.2009, 2 ♀♀, HG. Mirca, 28.iii.2010, 1 ♀, GS. Montaña de la Breña, 28.iii.2009, 2 ♀♀, BA. Pico Birigoyo, 9.vi.2011, 7 ♂♂ 7 ♀♀, GS. Puntagorda, 17.iii.2008, 1 ♀, JD (ZMAN). San Isidro, 6.v.2009, 1 ♂ 1 ♀, GS. Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS.

Taylorilygus apicalis (Fieber, 1861) Bco. Herradura, 12.vi.2011, 1 ♂, GS. Lomo Oscuro, 9.v.2009, 1 ♀, GS; 7.vi.2011, 3 ♀♀, GS. Puerto Naos, 24.iii.2010, 1 ♂ 1 ♀, CR, GS; 26.iii.2010, 1 ♂, CR. San Simón, 27.iii.2010, 2 ♀♀, CR.

Acetropis (Acetropis) gimmerthalii gimmerthalii (Flor, 1860) Fuencaliente N, 4.v.2009, 1 ♀, CR. Volcán Martín, 10.v.2009, 1 ♀, HG.

Dolichomiris (Dolichomiris) linearis Reuter, 1882 Bco. de las Angustias, 16.iv.2012, 1 ♂, HG. Jedey, 11.vi.2011, 1 ♂ 1 ♀, GS. Lomo Oscuro, 15.iii.2005, 1 ♂ 1 ♀, HG; 9.v.2009, 8 ♂♂ 10 ♀♀, HG, CR, GS; 20.iii.2010, 2 ♂♂ 3 ♀♀, CR, GS; 7.vi.2011, 1 ♂ 1 ♀, GS. Los Barros, 31.iii.2009, 1 ♂ 1 ♀, BA. Montaña Azufre, 27.iii.2010, 7 ♂♂ 5 ♀♀, CR, GS. Nogales, 6.–7.iii.2008, 2 ♂♂, JD (ZMAN). San Simón, 4.v.2009, 11 ♂♂ 8 ♀♀, CR, GS. Santo Domingo, 21.iii.2010, 1 ♀, GS.

466 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Heteroptera of La Palma, Canary Islands

Pachytomella passerinii (A. Costa, 1842) Montaña de la Breña, 28.iii.2009, 4 ♂♂ 2 ♀♀, BA. Montaña de la Negra, 25.iii.2010, 1 ♀, CR. Santo Domingo, 21.iii.2010, 1 ♂ 1 ♀, CR. El Palmar, 27.iii.2009, 7 ♂♂, BA.

Aetorhinella parviceps Noualhier, 1893 Bco. del Jorado, 10. and 13.vi.2011, 4 ♂♂ 11 ♀♀, GS. Bco. de la Galga, 1.v.2009, 6 ♂♂ 13 ♀♀, HG, CR, GS. El Pilar, 21.iii.2009, 1 ♀, BA. El Tablado, 9.iii.2008, 1 ♂, 15.iii.2008, 1 ♂, JD (ZMAN).

Canariocoris euphorbiae Lindberg, 1951 Lomo Oscuro, 9.v.2009, 7 ♂♂ 9 ♀♀, HG, CR, GS; 7.vi.2011, 1 ♂ 1 ♀, GS. Montaña Azufre, 27.iii.2010, 15 ♂♂ 9 ♀♀, on Euphorbia regis-jubae, CR, GS. Montaña la Laguna, 22.iii.2010, 1 ♀, GS.

Canariocoris pinicola (Lindberg, 1953) Bco. de las Angustias, 9.iii.2005, 1 ♂, HG. Fuencaliente N, 4.v.2009, 6 ♂♂ 3 ♀♀, CR, GS; 7.v.2009, 3 ♂♂, CR, GS; 10.v.2009, 2 ♂♂ 2 ♀♀, on Pinus canariensis, CR, GS. Jedey, 11.vi.2011, 8 ♂♂ 3 ♀♀, GS. Las Tricias, 8.v.2009, 6 ♂♂ 2 ♀♀, HG, CR, GS; 7.v.2011, 4 ♂♂ 5 ♀♀, GS. Los Barros, 31.iii.2009, 8 ♂♂ 4 ♀♀, BA. San Nicolás, 24.iii.2010, 1 ♂, GS. Volcán Martín, 10.v.2009, 1 ♂ 1 ♀, HG. Volcán San Antonio, 30.iii.2009, 1 ♂ 2 ♀♀, BA.

Canariocoris viburni (Lindberg, 1953) Bco. de la Galga, 1.v.2009, 1 ♂, CR; 28.iii.2010, 1 ♀, on Laurus, CR. El Tablado, 9.iii.2008, 1 ♀, JD (ZMAN).

Platycranus (Platycranus) lindbergi Wagner, 1954 Bco. Hondo, 20.iii.2010, 1 ♂ 2 ♀♀, CR. Fuencaliente N, 4.v.2009, 5 ♂♂ 8 ♀♀, CR, GS; 7.v.2009, 1 ♀, CR. Jedey, 11.vi.2011, 2 ♂♂ 1 ♀, GS. Las Tricias, 14.iii.2005, 1 ♂ 1 ♀, HG; 8.v.2009, 8 ♂♂ 8 ♀♀, on Retama, HG, CR, GS. Lomo Oscuro, 7.vi.2011, 2 ♀♀, GS. Los Llanos de Aridane, 20.iii.2009, 2 ♀♀, on Retama, BA. Montaña Azufre, 27.iii.2010, 1 ♂ 2 ♀♀, CR, GS. Montaña la Laguna, 13.vi.2007, 1 ♂ 3 ♀♀, HG; 2.v.2009, 18 ♂♂ 31 ♀♀, HG, CR, GS; 22.iii.2010, 2 ♂♂ 1 ♀, on Retama, CR. Puntagorda, 10.iii.2008, 9 ♂♂ 13 ♀♀, on Retama, JD (ZMAN). San Simón, 4.v.2009, 3 ♂♂ 4 ♀♀, CR, GS; 27.iii.2010 1 ♂ 1 ♀, on Retama, CR. Santo Domingo, 21.iii.2010, 3 ♂♂ 3 ♀♀, CR, GS.

Badezorus signaticornis (Reuter, 1904) El Pilar, 21.iii.2009, 1 ♀, BA. Puerto de Tazacorte, 14.iii.2008, 1 ♂ 4 ♀♀, JD (ZMAN).

Compsidolon (Coniortodes) beckeri (Reuter, 1904) Bco del Jorado, 23.iii.2009, 3 ♂♂ 4 ♀♀, BA. Bco. de las Angustias, 5.v.2009, 1 ♂, CR. Bco. Hondo, 30.iv.2009, 5 ♂♂ 1 ♀, CR, GS; 2.v.2009, 2 ♂♂ 1 ♀, on Adenocarpus, HG, CR. El Paso, farmland, 11.v.2009, 2 ♂♂ 2 ♀♀, GS. El Paso, gravel pit, 25.iii.2010, 1 ♂, CR; 27.iii.2010, 1 ♂ 1 ♀, on Chamaecytisus proliferus, CR. El Pilar, 17.vi.2007, 2 ♂♂ 1 ♀, HG; 3.v.2009, 2 ♂♂ 1 ♀, HG, CR, GS; 23.iii.2010, 1 ♀, CR, on Adenocarpus; 9.vi.2011, 6 ♂♂ 3 ♀♀, GS. Fuencaliente N, 4.v.2009, 1 ♂ 2 ♀♀, GS; 10.v.2009, 2 ♂♂ 1 ♀, HG. La Lomada, 18.vi.2007, 1 ♂, HG. Pico Birigoyo, 9.vi.2011, 6 ♂♂ 3 ♀♀, GS. Torre del Time, 1.vi.2011, 2 ♂♂ 1 ♀, GS. Volcán Martín, 10.v.2009, 9 ♂♂ 4 ♀♀, on Adenocarpus, CR, GS.

Compsidolon (Coniortodes) freyi (Wagner, 1954) Bco. del Jorado, 13.vi.2011, 1 ♂ 2 ♀♀, GS. Bco. de las Angustias, 20.iii.2005, 1 ♀, HG; 5.v.2009, 3 ♂♂ 7 ♀♀, HG, CR. Bco. Hondo, 30.iv.2009, 7 ♂♂ 11 ♀♀, HG, CR, GS; 2.v.2009, 5 ♂♂ 9 ♀♀, on Adenocarpus, HG,

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 467 B. AUKEMA ET AL.

CR. El Paso, farmland, 7.v.2009, 1 ♂, CR; 11.v.2009, 1 ♂, GS. El Paso, gravel pit, 5.vi.2011, 2 ♂♂ 2 ♀♀, GS. El Pilar, 17.vi.2007, 1 ♂ 1 ♀, HG; 9.vi.2011, 3 ♂♂ 2 ♀♀, GS. Fuencaliente, 9.vi.2007, 7 ♂♂ 10 ♀♀, HG. Fuencaliente N, 4.v.2009, 1 ♂, CR. Jedey, 11.vi.2011, 1 ♂ 1 ♀, GS. Lomo Oscuro, 15.iii.2005, 1 ♂ 1 ♀, HG. Los Llanos de Aridane, 20.iii.2009, 1 ♂ 1 ♀, BA. Montaña la Laguna, 2.v.2009, 1 ♀, on Retama, CR. Nogales, 6.–7.iii.2008, 1 ♂, JD (ZMAN). Pico Birigoyo, 9.vi.2011, 3 ♂♂ 2 ♀♀, GS. Puntagorda, 15.iii.2008, 1 ♀, JD (ZMAN). Roque de los Muchachos, 27.ii.2001, 2 ♂♂ 2 ♀♀, HG. Salinas, 19.vi.2007, 1 ♂, HG. San Isidro, 6.v.2009, 1 ♂, HG. San Simón, 4.v.2009, 2 ♂♂, CR, GS. Torre del Time, 1.vi.2011, 4 ♂♂ 3 ♀♀, GS. Volcán Martín, 21.ii.2001, 1 ♂ 4 ♀♀, HG; 10.v.2009, 15 ♂♂ 20 ♀♀, on Adenocarpus, GS. Volcán Teneguía, 11.vi.2007, 1 ♂, HG.

Compsidolon (Coniortodes) hierroense (Wagner, 1954) Bco. de las Angustias, 9.iii.2005, 5 ♂♂ 3 ♀♀; 5.v.2009, 1 ♀, HG, GS. Bco. Hondo, 2.v.2009, 2 ♂♂ 1 ♀, HG. El Palmar, 27.iii.2009, 2 ♂♂ 1 ♀, BA. El Paso, farmland, 11.vi.2007, 1 ♀, GS; 11.v.2009, 1 ♀. El Paso, gravel pit, 5.vi.2011, 2 ♂♂ 1 ♀, GS. Fuencaliente N, 4.v.2009, 1 ♂ 2 ♀♀, CR, GS; 10.v.2009, 3 ♂♂ 4 ♀♀, on Sideritis, CR. Laguna de Barlovento, 21.iv.2012, 1 ♂ 1 ♀, HG. Las Tricias, 8.v.2009, 1 ♂ 4 ♀♀, GS. Lomo Oscuro, 9.vi.2007, 1 ♀, HG. Puntagorda, 17.iii.2008, 2 ♀♀. Tijarafe, 12.iii.2008, 3 ♂♂ 1 ♀, JD (ZMAN). Torre del Time, 1.vi.2011, 2 ♂♂ 6 ♀♀, GS.

Compsidolon (Coniortodes) longiceps (Reuter, 1904) Bco. de las Angustias, 5.v.2009, 3 ♂♂ 4 ♀♀, CR, GS. Los Barros, 31.iii.2009, 1 ♂ 3 ♀♀, BA.

Compsidolon (Coniortodes) verbenae (Wagner, 1954) Bco. Herradura, 12.vi.2011, 1 ♂ 3 ♀♀, GS. Fuencaliente NE, 4.v.2009, 6 ♂♂ 15 ♀♀, on Artemisia, CR, GS. La Lomadita, 21.iii.2010, 1 ♂ 1 ♀, CR, GS. Lomo Oscuro, 15.iii.2005, 1 ♂ 2 ♀♀, HG; 9.v.2009, 23 ♂♂ 22 ♀♀, CR, GS; 20.iii.2010, 2 ♂♂ 1 ♀, on Artemisia, GS. Montaña Azufre, 27.iii.2010, 1 ♂, GS. Puntagorda, 17.iii.2008, 4 ♂♂ 6 ♀♀, on Artemisia, JD (ZMAN). Volcán Martín, 21.ii.2001, 3 ♂♂, HG.

Conostethus venustus longicornis (Fieber, 1858) El Paso, gravel pit, 23.iii.2010, 3 ♀♀, CR; 25.iii.2010, 1 ♀, CR; 27.iii.2010, 3 ♂♂ 8 ♀♀, GS. Las Tricias, 14.iii.2005, 3 ♀♀. Montaña de la Negra, 25.iii.2010, 1 ♂ 2 ♀♀, CR. Santo Domingo, 21.iii.2010, 17 ♂♂ 18 ♀♀, CR, GS; 26.iii.2010, 6 ♂♂ 1 ♀, CR.

Lindbergopsallus hyperici (Lindberg, 1953) Bco. de las Angustias, 3.vi.2011, 9 ♂♂ 8 ♀♀. Callejones, 27.iii.2010, 1 ♂ 1 ♀, on Hypericum, CR. La Lomada, 18.vi.2007, 2 ♂♂ 1 ♀, HG. Lomo Oscuro, 7.vi.2011, 2 ♂♂ 1 ♀. Mirca, 28.iii.2010, 35 ♂♂ 12 ♀♀, on Hypericum canariense, CR. Montaña de la Breña, 28.iii.2009, 4 ♂♂ 2 ♀♀, on Hypericum, BA.

Lindbergopsallus instabilis (Reuter, 1904) Bco. del Jorado, 23.iii.2009, 2 ♂♂ 3 ♀♀, BA. Bco. de las Angustias, 5.v.2009, 1 ♂, CR. La Lomadita, 21.iii.2010, 7 ♂♂ 6 ♀♀, CR, GS. Las Tricias, 8.v.2009, 1 ♂, HG. Lomo Oscuro, 15.iii.2005, 2 ♂♂ 2 ♀♀, HG; 9.v.2009, 2 ♂♂ 2 ♀♀, HG, CR; 20.iii.2010, 15 ♂♂ 19 ♀♀, CR, GS; 7.vi.2011, 3 ♀♀, on Lavendula, GS. Los Llanos de Aridane, 20.iii.2009, 2 ♀♀, on Lavendula, BA. Montaña Azufre, 27.iii.2010, 15 ♂♂ 12 ♀♀, CR, GS. Montaña la Laguna, 2.v.2009, 2 ♂♂, HG, CR, 22.iii.2010, 4 ♂♂ 10 ♀♀, on Lavendula, CR, GS. Nogales, 6–7.iii.2008, 8 ♂♂ 2 ♀♀, JD (ZMAN). Puerto Naos, 24.iii.2010, 1 ♂ 4 ♀♀, GS; 26.iii.2010, 1 ♀, on Lavendula, CR. Punta Baja de San Simón, 4.iv.2009, 2 ♂♂, BA. Puntagorda, 10.iii.2008, 1 ♀, JD (ZMAN). San Simón, 27.iii.2010, 1 ♀, CR. Santo Domingo, 21.iii.2010, 9 ♂♂ 5 ♀♀, GS.

468 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Heteroptera of La Palma, Canary Islands

Lindbergopsallus lunariae Rieger, Strauss et Günther, 2011 Bco. del Jorado, 23.iii.2009, 4 ♂♂ 4 ♀♀, BA. Bco. de las Angustias, 20.iii.2005, 1 ♂, HG; 16.iv.2012, 3 ♂♂ 1 ♀, HG. El Paso, farmland, 24.iv.2012, 2 ♂♂ 1 ♀, HG. Fuencaliente N, 4.v.2009, 6 ♂♂, CR; 7.v.2009, 2 ♀♀, CR; 10.v.2009, 2 ♂♂ 1 ♀, HG. Fuencaliente NE, 4.v.2009, 3 ♂♂ 2 ♀♀, CR, GS. La Fajana, 28.ii.1998, 7 ♂♂, on Rumex lunaria, JD (ZMAN). Lomo Oscuro, 15.iii.2005, 2 ♂♂, HG; 9.vi.2007, 1 ♂, HG; 9.v.2009, 13 ♂♂ 3 ♀♀, HG, GS; 20.iii.2010, 6 ♂♂ 8 ♀♀, CR, GS; 7.vi.2011, 3 ♂♂ 2 ♀♀, GS; 14.iv.2012, 1 ♂. Montaña Argual, 29.iii.2009, 7 ♂♂ 2 ♀♀, BA. Montaña Azufre, 27.iii.2010, 9 ♂♂ 1 ♀. Montaña la Laguna, 2.v.2009, 1 ♂ 2 ♀♀, CR, GS. Puerto de Tazacorte, 14.iii.2008, 2 ♂♂ 1 ♀. Puntagorda 15.iii.2008, 7 ♂♂ 12 ♀♀, JD (ZMAN). Torre del Time, 1.vi.2011, 1 ♂ 2 ♀♀, GS. Volcán Teneguía, 11.vi.2007, 1 ♀, HG; 30.iv.2009, 1 ♂, GS; 20.iv.2012, 3 ♂♂ 1 ♀, HG.

Lindbergopsallus lauri Rieger, 2012 Montaña de la Breña, 28.iii.2009, 1 ♂, BA.

Tuponia (Chlorotuponia) oculata Wagner, 1954 Bco. Herradura, 12.vi.2011, 1 ♂ 8 ♀♀, GS. Punta Baja de San Simón, 4.v.2009, 5 ♂♂ 4 ♀♀, on Tamarix, CR, GS.

Tuponia (Tuponia) rubella Puton, 1889 Bco. de la Galga, 24.ii.2001, 2 ♂♂, HG. El Pilar, 21.iii.2009, 1 ♂, BA. La Lomada, 18.vi.2007, 1 ♂, HG. Lomo Oscuro, 9.vi.2007, 2 ♂♂, HG. Puntagorda, 10.iii.2008, 1 ♂ 1 ♀. El Tablado, 9.iii.2008, 1 ♂, JD (ZMAN).

NABIDAE Nabis (Nabis) pseudoferus ibericus Remane, 1962 Bco. de la Galga, 12.vi.2007, 3 ♂♂ 2 ♀♀, HG; 1.v.2009, 4 ♂♂ 3 ♀♀, HG, CR, GS; 28.iii.2010, 2 ♀♀, CR. El Paso, farmland, 11.vi.2007, 1 ♂ 1 ♀, HG; 7.v.2009, 1 ♀; 11.v.2009, 2 ♂♂, HG, CR; 22.iii.2010, 2 ♀♀, CR. El Pilar, 17.vi.2007, 1 ♂, HG. La Lomada, 18.vi.2007, 1 ♂ 1 ♀, HG; Laguna de Barlovento, 1.iv.2009, 1 ♀, BA. Lomo Oscuro, 9.vi.2007, 1 ♂ 1 ♀, HG; 9.v.2009, 1 ♀, HG. Montaña Azufre, 4.v.2009, 1 ♂, GS. Roque de los Muchachos, 27.ii.2001, 1 ♀, HG. San Isidro, 6.v.2009, 5 ♂♂ 1 ♀, CR. San Simón, 4.v.2009, 1 ♂ 1 ♀, GS; 27.iii.2010, 2 ♀♀, CR.

Nabis (Tropiconabis) capsiformis Germar, 1838 Bco. de la Galga, 1.v.2009, 1 ♂, GS. Bco. de las Angustias, 9.iii.2005, 1 ♀, HG; 20.iii.2005, 2 ♂♂ 3 ♀♀, HG; 3.vi.2011, 1 ♂ 1 ♀, GS. Bco. Hondo, 2.v.2009, 1 ♀, GS. El Paso, farmland, 11.v.2009, 1 ♂, GS. Fuencaliente, 26.ii.2001, 1 ♀, HG. Las Tricias, 7.v.2011, 1 ♂ 2 ♀♀, GS. Lomo Oscuro, 9.v.2009, 1 ♂ 3 ♀♀, HG, GS; 20.iii.2010, 1 ♂, GS. Montaña Azufre, 27.iii.2010, 1 ♂ 1 ♀, CR, GS. Montaña la Laguna, 2.v.2009, 1 ♀, GS; 22.iii.2010, 1 ♀, GS. Puerto Naos, 24.iii.2010, 1 ♂, GS. San Simón, 4.v.2009, 2 ♂♂ 2 ♀♀, CR. Torre del Time, 1.vi.2011, 1 ♂, GS.

ANTHOCORIDAE Anthocoris alienus (White, 1880) Bco. de las Angustias, 5.v.2009, 1 ♂, HG. Bco. Hondo, 2.v.2009, 2 ♂♂, CR. El Paso, gravel pit, 25.iii.2010, 4 ♂♂ 3 ♀♀, CR; 27.iii.2010, 3 ♂♂ 10 ♀♀ on Chamaecytisus proliferus, CR. El Pilar, 17.vi.2007, 1 ♀, HG;

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3.v.2009, 18 ♂♂ 38 ♀♀, on Adenocarpus, HG, CR, GS. Pico Birigoyo, 9.vi.2011, 3 ♂♂ 4 ♀♀, GS. Roque de los Muchachos, 26.ii.2001, 2 ♀♀, HG; 27.ii.2001, 1 ♂ 1 ♀, HG. Volcán Martín, 10.v.2009, 1 ♀ 1 larva, on Adenocarpus, CR.

Anthocoris salicis Lindberg, 1953 Barlovento, 21. and 26.v.2010, 3 ♀♀ 10 ♀♀, on Laurus, CR. Bco. de la Galga, 28.iii.2010, 1 ♂ 1 ♀, on Laurus, GS. El Paso, gravel pit, 27.iii.2010, 1 ♀, on Chamaecytisus, CR. El Pilar, 21.iii.2009, 1 ♀, BA. Santa Cruz, 16.iii.2005, 1 ♀, HG.

Orius (Microtrachelia) retamae (Noualhier, 1893) Roque de los Muchachos, 27.ii.2001, 3 ♀♀, HG; 8.iii.2008, 1 ♂ 14 ♀♀, on Adenocarpus spartioides, JD (ZMAN).

Orius (Orius) laevigatus maderensis (Reuter, 1884) Bco. de la Galga, 1.v.2009, 2 ♀♀, HG, CR. Bco. de las Angustias, 5.vi.2007, 1 ♂, HG; 5.v.2009, 1 ♀, HG. Bco. Hondo, 30.iv.2009, 3 ♂♂ 10 ♀♀, CR, GS; 2.v.2009, 1 ♂ 11 ♀♀, HG, CR. El Paso, farmland, 7.v.2009, 1 ♀, CR. El Pilar, 2.v.2009, 2 ♂♂ 16 ♀♀, HG, GS. Fuencaliente N, 4.v.2009, 2 ♀♀, on Rumex, CR; 10.v.2009, 2 ♀♀, HG. Fuencaliente NE, 4.v.2009, 1 ♂ 1 ♀, on Artemisia, CR. La Lomadita, 21.iii.2010, 10 ♂♂ 7 ♀♀, GS. Las Tricias, 8.v.2009, 1 ♀, CR. Lomo Oscuro, 9.vi.2007, 1 ♂ 1 ♀, HG; 9.v.2009, 2 ♂♂ 2 ♀♀, HG, CR. Montaña Azufre, 27.iii.2010, 1 ♂, GS. Montaña la Laguna, 12.vi.2007, 1 ♂, HG; 2.v.2009, 1 ♂, CR; 22.iii.2010, 3 ♂♂ 1 ♀, CR. Nogales, 6–7.iii.2008, 2 ♂♂ 3 ♀♀, J.D. (ZMAN). Pico de la Nieve, 6.v.2009, 1 ♀, HG. Roque de los Muchachos, 27.ii.2001, 1 ♂ 1 ♀, HG. Volcán Martín, 10.v.2009, 7 ♂♂ 6 ♀♀, on Adenocarpus, CR. Volcán San Antonio, 30.iii.2009, 1 ♂, BA. Volcán Teneguía, 30.iv.2009, 5 ♂♂ 5 ♀♀, CR.

Orius (Orius) limbatus Wagner, 1952 La Lomadita, 21.iii.2010, 28 ♂♂ 15 ♀♀, CR. Los Llanos de Aridane, 26.iii.2009, 1 ♂, BA. Volcán San Antonio, 30.iii.2009, 2 ♂♂ 2 ♀♀, BA.

Xylocoris (Xylocoris) contiguus Wagner, 1954 Bco. Hondo, 2.v.2009, 1 larva, HG. Fuencaliente N, 7.v.2009, 3 ♂♂, in dried grass tufts, CR, GS; 10.v.2009, 1 ♂, HG. Lomo Oscuro, 9.v.2009, 3 ♀♀, in dried grass tufts, CR. Volcán Martín, 10.v.2009, 2 ♂♂ 1 ♀, HG.

REDUVIIDAE rubromaculatus (Blackburn, 1889) Bco. de las Angustias, 5.v.2009, 1 ♀, CR. Puerto Naos, 26.iii.2010, 2 ♂♂ 2 ♀♀, on Nicotiana glauca, CR.

Coranus (Coranus) griseus (Rossi, 1790) Lomo Oscuro, 20.iii.2010, 1 larva 5 → 1 ♀, CR. Montaña la Laguna, 22.iii.2010, 1 ♀, GS. Puerto Naos, 24.iii.2010, 2 ♀♀ 1 larva 5 → 1 ♂, CR, GS; 29.iii.2010, 1 ♂ 3 ♀♀ 3 larvae 5 ? 2 ♂♂ 1 ♀, CR, GS. Las Tricias, 7.v.2011, 1 ♂, GS.

Coranus (Coranus) aegyptius (Fabricius, 1775) El Paso, farmland, 7.v.2009, 1 ♂ 1 ♀, HG. Lomo Oscuro, 9.v.2009, 1 ♂, HG. Montaña la Laguna, 12.vi.2007, 3 ♂♂, HG.

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ARADIDAE Aneurus (Aneurodes) tagasastei Enderlein, 1931 Bco. del Agua, 29.ix.2002, 2 ♂♂ 3 ♀♀, W. Löderbusch leg. (CR). Bco. Gallegos, 5.x.2002, 4 ♂♂ 2 ♀♀, F. Lange leg. (CR). Breña Alta, 1.x.2002, 1 ♂, F. Lange leg. (CR). El Tablado, 9.iii.2008, 1 ♀ 1 larva 5, JD (ZMAN).

Aradus (Aradus) reuterianus Puton, 1875 Hoya Grande, 30.ix.2002, 1 ♀, F. Lange leg. (CR).

LYGAEIDAE Caenocoris nerii (Germar, 1847) Puntagorda, 10.iii.2008, 1 ♂, JD (ZMAN). Volcán Teneguía, 11.vi.2007, 1 ♂, HG.

Lygaeosoma sardeum sardeum Spinola, 1837 Bco. Hondo, 30.iv.2009, 1 ♀, GS. El Palmar, 27.iii.2009, 1 ♀, BA. La Punta, 2.iv.2009, 1 ♀, BA. Las Tricias, 7.v.2011, 1 ♂ 1 ♀, GS. Nogales, 6–7.iii.2008, 1 ♀, JD (ZMAN). Puntagorda, 17.iii.2008, 1 ♀, JD (ZMAN); 25.iii.2010, 1 ♂, GS. San Simón, 27.iii.2010, 3 ♂♂ 2 ♀♀, CR. Santo Domingo, 21.iii.2010, 10 ♂♂ 2 ♀♀, CR, GS; 26.iii.2010, 2 ♂♂ 3 ♀♀, CR.

Spilostethus pandurus (Scopoli, 1763) Bco. de las Angustias, 3.vi.2011, 1 ♂, GS. Bco. Hondo, 30.iv.2009, 1 ♀, GS. El Paso, farmland, 7.v.2009, 1 ♀, CR. Fuencaliente N, 20.iii.2010, 1 ♂, CR. Jedey, 13.iii.2008, 1 ♀, JD (ZMAN). La Fajana, 9.iii.2008, 1 ♀, JD (ZMAN). La Lomadita, 21.iii.2010, 1 ♂, GS. Las Tricias, 8.v.2009, 1 ♀, GS. Puntagorda, 23.iii.2010, 1 ♀, GS. Torre del Time, 1.vi.2011, 1 ♀, GS.

Nysius cymoides (Spinola, 1837) El Pilar, 21.iii.2009, 1 ♀, BA. Jedey, 11.vi.2011, 3 ♂♂ 3 ♀♀, GS. Lomo Oscuro, 7.vi.2011, 3 ♂♂ 1 ♀, GS.

Nysius ericae ericae (Schilling, 1829) Fuencaliente, 26.ii.2001, 1 ♀, HG. Lomo Oscuro, 9.vi.2007, 1 ♂ 2 ♀♀, HG; 9.v.2009, 1 ♂, GS. Roque de los Muchachos, 27.ii.2001, 1 ♀, HG; 13.vi.2007, 1 ♂, HG. Salinas, 12.vi.2007, 1 ♂ 1 ♀, HG. San Simón, 9.vi.2007, 1 ♀; 4.v.2009, 1 ♂ 1 ♀, GS. Torre del Time, 1.vi.2011, 1 ♀, GS.

Nysius immunis (Walker, 1872) Barlovento, 21.iii.2010, 2 ♂♂ 2 ♀♀, GS. Bco. del Jorado, 10.vi.2011, 1 ♀, GS; 13.vi.2011, 1 ♂ 1 ♀, GS. Bco. de las Angustias, 21.ii.2001, 1 ♂, HG; 3.vi.2011, 1 ♂ 1 ♀, GS. Bco. Herradura, 12.vi.2011, 1 ♂ 1 ♀, GS. Callejones, 27.iii.2010, 1 ♂, on Hypericum, CR. El Pilar, 17.vi.2007, 1 ♂ 1 ♀, HG; 3.v.2009, 1 ♂ 1 ♀, HG; 23.iii.2010, 1 ♀, GS. Fuencaliente N, 7.v.2009, 4 ♂♂ 2 ♀♀, CR, GS. Fuencaliente NE, 4.v.2009, 1 ♂ 1 ♀, CR. Jedey S, 11.vi.2011, 2 ♂♂ 2 ♀♀, GS. La Fajana, 9.iii.2008, 1 ♂ 1 ♀, JD (ZMAN). La Lomadita, 21.iii.2010, 1 ♂ 1 ♀, CR, GS. Lomo Oscuro, 9.v.2009, 2 ♂♂ 6 ♀♀, HG, GS; 20.iii.2010, 2 ♂♂ 2 ♀♀, CR, GS; 7.vi.2011, 3 ♂♂ 1 ♀, GS. Montaña de la Negra, 25.iii.2010, 1 ♂ 1 ♀, CR. Montaña la Laguna, 13.vi.2007, 1 ♀, HG; 2.v.2009, 8 ♂♂ 8 ♀♀, HG, CR, GS. Nogales, 6–7.iii.2008, 4 ♂♂, JD (ZMAN). Puntagorda, 25.iii.2010, 1 ♂, from leaf crown of Sonchus, CR, GS. Puntagorda, 17.iii.2008, 2 ♂♂, JD (ZMAN). Roque de los Muchachos, 27.ii.2001, 1 ♀, HG; 15.iii.2005, 1 ♂ 1 ♀, HG. Salinas, 12.vi.2007, 1 ♂, HG. San Isidro, 6.v.2009, 1 ♀, GS. San Nicolás, 24.iii.2010, 2 ♂♂ 2 ♀♀, CR, GS. Santo Domingo, 21.iii.2010, 1 ♂, CR. Volcán Teneguía, 26.ii.2001, 2 ♀♀, HG; 30.iv.2009, 4 ♂♂ 2 ♀♀, HG, CR.

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Orsillus pinicanariensis Lindberg, 1953 Bco. de las Angustias, 5.v.2009, 2 ♀♀, on Pinus, CR. El Paso, gravel pit, 23.iii.2010, 1 ♂, CR; 27.iii.2010, 1 ♀, GS; 5.vi.2011, 1 ♂, GS. El Pilar, 2.v.2009, 1 ♂, GS; 23.iii.2010, 3 ♂♂ 2 ♀♀, on Pinus canariensis, CR. El Pinar, 25.iii.2009, 4 ♂♂, on Pinus canariensis, BA. Fuencaliente N, 4.v.2009, 2 ♂♂ 2 ♀♀, on Pinus, CR, GS. Las Tricias, 7.v.2011, 1 ♂, GS. San Nicolás, 20.iii.2010, 4 ♂♂ 20 ♀♀, CR, GS; 24.iii.2010, 1 ♂ 3 ♀♀, in pine cones, GS. Torre del Time, 1.vi.2011, 1 ♂, GS. Volcán Martín, 10.v.2009, 5 ♂♂ 2 ♀♀, on Pinus, HG, CR, GS.

Kleidocerys truncatulus (Walker, 1872) Barlovento, 21.iii.2010, 2 ♂♂ 2 ♀♀, on Erica, CR. Bco. de la Galga, 24.ii.2001, 1 ♂, HG; 1.v.2009, 10 ♂♂ 6 ♀♀, HG, on Erica arborea, CR; 28.iii.2010, 1 ♂, GS. El Paso, farmland, 7.v.2009, 1 ♀, HG; 11.v.2009, 1 ♂ 1 ♀, on Erica arborea, CR; 22.iii.2010, 1 ♂, GS. El Pilar, 21.iii.2009, 2 ♂♂ 1 ♀, BA; 3.v.2009, 3 ♂♂ 4 ♀♀, on Erica arborea, CR, GS; 23.iii.2010, 1 ♂ 1 ♀, on Erica, CR, GS; 9.vi.2011, 1 ♂ 3 ♀♀, GS. El Tablado, 9.iii.2008, 2 ♂♂ 1 ♀; 15.iii.2008, 2 ♂♂, JD (ZMAN). Laguna de Barlovento, 1.iv.2009, 1 ♂ 2 ♀♀, on Erica arborea, BA. Lomo Oscuro, 9.v.2009, 1 ♂, HG. Pico Birigoyo, 9.vi.2011, 1 ♂ 3 ♀♀, GS. San Isidro, 6.v.2009, 4 ♂♂ 1 ♀, HG, GS; 23.iv.2012, 1 ♂, HG. Santo Domingo, 21.iii.2010, 2 ♂♂ 1 ♀, CR, GS.

Geocoris (Geocoris) pubescens (Jakovlev, 1871) Bco. de la Galga, 12.vi.2007, 1 ♂, HG. El Paso, 11.vi.2007, 1 ♀, HG. Hoya Grande, 8.iii.2008, 1 ♂ 1 ♀, JD (ZMAN). Lomo Oscuro, 9.v.2009, 1 ♂, HG. Puerto de Tazacorte, 14.iii.2008, 1 ♀, JD (ZMAN).

Heterogaster canariensis Lindberg, 1960 Bco. del Jorado, 13.vi.2011, 2 ♂♂, GS. Bco. de las Angustias, 22.ii.2001, 2 ♂♂ 1 ♀, HG; 16.iv.2012, 1 ♂ 1 ♀, HG. Bco. Hondo, 30.iv.2009, 1 ♀, GS. El Paso, farmland, 11.v.2009, 4 ♂♂, CR. Jedey, 11.vi.2011, 2 ♂♂ 2 ♀♀, GS. La Lomadita, 21.iii.2010, 1 ♂ 1 ♀, GS. Las Tricias, 8.v.2009, 8 ♂♂ 3 ♀♀, CR, GS; 7.vi.2011, 1 ♀, GS. Lomo Oscuro, 7 ♂♂ 3 ♀♀, HG; 9.v.2009, 1 ♂ 1 ♀, on Thymus, CR; 20.iii.2010, 2 ♂♂, on Lavendula, CR, GS; 7.vi.2011, 1 ♂ 1 ♀, GS. Los Llanos de Aridane, 20.iii.2009, 3 ♀♀, on Lavendula, BA. Montaña Azufre, 27.iii.2010, 1 ♂ 1 ♀, GS. Montaña la Laguna, 2.v.2009, 5 ♂♂ 4 ♀♀, on Lavendula, CR, GS; 22.iii.2010, 4 ♂♂ 3 ♀♀, CR, GS. Puerto Naos, 24.iii.2010, 1 ♀, GS; 29.iii.2010, 1 ♀, on Lavendula, CR. Punta Baja de San Simón, 4.v.2009, 1 ♂ 6 ♀♀, on Lavendula, CR, GS. San Nicolás, 24.iii.2010, 2 ♂♂ 2 ♀♀, CR, GS. Santo Domingo, 21.iii.2010, 1 ♂ 3 ♀♀, GS. Volcán Martín, 21.ii.2001, 2 ♂♂ 1 ♀, HG. Volcán San Antonio, 30.iii.2009, 2 ♀♀, on Lavendula, BA.

Heterogaster urticae (Fabricius, 1775) Punta Baja de San Simón, 20.iii.2010, 4 larvae → 2 ♀♀, CR; 27.iii.2010, 6 larvae → 1 ♂ 5 ♀♀, CR.

Bethylimorphus leucophaes Lindberg, 1953 Las Tricias, 7.v.2011, 1 ♂, GS. Lomo Oscuro, 15.iii.2005, 2 ♂♂ 4 ♀♀; 4.v.2009, 2 ♂♂ 1 ♀, GS. Montaña Azufre, 27.iii.2010, 1 ♂, CR. Montaña la Laguna, 12.vi.2007, 2 ♂♂ 2 ♀♀. Salinas, 6.xii.2007, 4 ♂♂ 2 ♀♀. San Simón, 4.v.2009, 2 ♂♂ 1 ♀.

Macroplax vicina Puton, 1889 Bco. del Jorado, 13.vi.2011, 1 ♂, GS. Bco. Hondo, 30.iv.2009, 4 ♂♂ 3 ♀♀, CR, GS; 2.v.2009, 10 ♂♂ 5 ♀♀, HG, CR. El Paso, farmland, 11.vi.2007, 2 ♂♂ 1 ♀, HG; 7.v.2009, 4 ♂♂ 1 ♀, HG, CR; 11.v.2009, 4 ♂♂ 4 ♀♀, CR, GS; 22.iii.2010, 5 ♂♂ 4 ♀♀, CR, GS. El Paso, gravel pit, 25.iii.2010, 1 ♂, CR; 27.iii.2010, 1 ♀, GS. El Pilar, 17.vi.2007, 13 ♂♂ 9 ♀♀, HG; 21.iii.2009, 2 ♂♂ 4 ♀♀, BA; 3.v.2009, 5 ♂♂ 5 ♀♀, CR, GS; 23.iii.2010, 1 ♂ 1 ♀, GS. El Pinar, 25.iii.2009, 6 ♂♂ 6 ♀♀, BA. Fuencaliente N, 7.v.2009, 1 ♀, GS. Jedey, 11.vi.2011, 1 ♂ 1 ♀, GS. Las Tricias, 8.v.2009, 2 ♂♂ 3 ♀♀, CR, GS. Lomo Oscuro, 9.v.2009, 1 ♂, CR. Pico Birigoyo, 9.vi.2011, 1 ♂ 1 ♀, GS. Pico de la Nieve, 1070 m a.s.l., 6.v.2009, 5 ♂♂ 3 ♀♀, on Cistus symphytifolius, CR. Pico de la Nieve, 665 m a.s.l., 6.v.2009, 3 ♂♂ 2 ♀♀, on Cistus monspeliensis, HG, CR. Puntagorda, 25.iii.2010,

472 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Heteroptera of La Palma, Canary Islands

1 ♂, GS. San Isidro, 6.v.2009, 5 ♂♂ 5 ♀♀, HG, CR. Santo Domingo, 21.iii.2010, 1 ♂, GS. Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS.

Microplax interrupta (Fieber, 1837) Bco. Hondo, 30.iv.2009, 2 ♂♂ 4 ♀♀, HG, CR, GS; 2.v.2009, 6 ♂♂ 3 ♀♀, HG, CR. El Pilar, 3.v.2009, 3 ♂♂ 2 ♀♀, CR, GS. Garafia, 17.iii.2008, 6 ♂♂ 6 ♀♀, JD (ZMAN). San Simón, 27.iii.2010, 8 ♂♂ 13 ♀♀, CR. Santo Domingo, 21.iii.2010, 1 ♂ 1 ♀, GS.

Oxycarenus (Oxycarenus) lavaterae (Fabricius, 1787) Bco. del Jorado, 10.vi.2011, 1 ♂ 1 ♀, GS. Bco. de las Angustias, 3.vi.2011, 1 ♂, GS. Bco. Herradura, 12.vi.2011, 2 ♀♀, GS. Bco. Hondo, 30.iv.2009, 1 ♀, GS. Jedey, 13.iii.2008, 1 ♀, JD (ZMAN). La Lomadita, 21.iii.2010, 4 ♂♂ 4 ♀♀, CR, GS. La Punta, 2.iv.2009, 1 ♂, BA. Las Tricias, 8.v.2009, 2 ♂♂ 3 ♀♀, on cultivated Malva, CR, GS. Pico de la Nieve, 665 m a.s.l., 6.v.2009, 1 ♂, GS. Puerto Naos, 24.iii.2010, 3 ♂♂ 4 ♀♀, CR, GS; 29.iii.2010, 1 ♂ 1 ♀, GS. Puntagorda, 25.iii.2010, 1 ♀, CR. Tazacorte, 21.iii.2010, 1 ♂, GS. Torre del Time, 1.vi.2011, 1 ♀, GS.

Tropistethus seminitens Puton, 1889 Bco. de la Galga, 24.ii.2001, 1 ♀, HG. Bco. de las Angustias, 20.iii.2005, 1 ♂ 1 ♀, HG. Bco. Hondo, 2.v.2009, 1 ♂, GS. El Pilar, 3.v.2009, 5 ♂♂ 6 ♀♀, CR, GS. Fuencaliente N, 20.iii.2010, 24 ♂♂ 17 ♀♀, CR, GS. Laguna de Barlovento, 21.iv.2012, 1 ♂ 1 ♀, HG. Lomo Oscuro, 15.iii.2005, 3 ♂♂ 4 ♀♀, HG; 9.v.2009, 1 ♂, CR; 20.iii.2010, 1 ♀, GS. Los Llanos de Aridane, 26.iii.2009, 5 ♂♂, BA. San Isidro, 6.v.2009, 2 ♀♀, GS. Volcán Martín, 21.ii.2001, 1 ♂ 1 ♀, HG.

Eremocoris maderensis (Wollaston, 1858) San Isidro, 6.v.2009, 1 ♀, HG.

Ischnocoris mundus (Walker, 1872) Bco. Hondo, 30.iv.2009, 1 ♀, GS. El Paso, farmland, 7.v.2009, 1 ♂, CR. El Pilar, 2.v.2009, 1 ♂ 2 ♀♀, GS. El Pinar, 25.iii.2009, 2 ♀♀, BA. San Isidro, 6.v.2009, 1 ♂ 2 ♀♀, CR, GS.

Scolopostethus pilosus maderensis Reuter, 1881 Bco. de la Galga, 24.ii.2001, 2 ♂♂ 2 ♀♀, HG; 10.iii.2005, 1 ♂ 1 ♀, HG; 1.v.2009, 2 ♀♀, HG, GS. El Pilar, 3.v.2009, 2 ♂♂ 4 ♀♀, CR. El Tablado, 9.iii.2008, 5 ♂♂ 6 ♀♀; 15.iii.2008, 5 ♀♀ 1 larva 5, JD (ZMAN). Fuencaliente, 26.ii.2001, 1 ♂, HG. Laguna de Barlovento, 1.iv.2009, 4 ♂♂ 4 ♀♀, BA; 21.iv.2012, 2 ♂♂ 2 ♀♀, HG. Montaña la Laguna, 12.vi.2001, 1 ♀, HG. San Isidro, 6.v.2009, 8 ♂♂ 6 ♀♀, HG, CR, GS.

Aphanus rolandri (Linnaeus, 1758) Bco. de la Galga, 10.iii.2005, 2 ♂♂, HG. El Paso, farmland, 22.iii.2010, 1 ♂, CR. Garafia, 11.iii.2008, 1 ♂, JD (ZMAN). Laguna de Barlovento, 1.iv.2009, 4 ♂♂ 1 ♀, BA. Los Llanos de Aridane, 23.iii.2009, 1 ♂; 26.iii.2009, 1 ♀, BA. Volcán Martín, 21.ii.2001, 1 ♂ 1 ♀, HG.

Emblethis angustus Montandon, 1890 El Palmar, 27.iii.2009, 1 ♂, BA; Laguna de Barlovento, 21.iv.2012, 1 ♀, HG.

Emblethis denticollis Horváth, 1878 El Palmar, 27.iii.2009, 2 ♂♂ 2 ♀♀, BA. El Paso, farmland, 7.v.2009, 1 ♂ 1 ♀, CR. El Pilar, 21.iii.2009, 1 ♀, BA. La Punta, 2.iv.2009, 6 ♂♂ 6 ♀♀, BA.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 473 B. AUKEMA ET AL.

Emblethis duplicatus Seidenstücker, 1963 El Palmar, 27.iii.2009, 3 ♀♀, BA. El Paso, farmland, 7.v.2009, 1 ♂; 22.iii.2010, 1 ♂ 2 ♀♀, CR, GS. El Paso, gravel pit, 25.iii.2010, 1 ♂ 3 ♀♀, CR; 27.iii.2010, 1 ♂ 3 ♀♀, GS; 5.vi.2011, 1 ♂ 1 ♀, GS. La Lomadita, 23.iv.2012, 1 ♀. Laguna de Barlovento, 21.iv.2012, 2 ♂♂ 1 ♀, HG. Lomo Oscuro, 23.iv.2012, 1 ♀, HG. Los Llanos de Aridane, 26.iii.2009, 1 ♀, BA. Nogales, 6–7.iii.2008, 1 ♀, JD (ZMAN). Puerto Naos, 29.iii.2010, 1 ♂, GS. San Simón, 9.vi.2007, 1 ♀, 27.iii.2010, 1 ♂, CR. Santo Domingo, 21.iii.2010, 1 ♀, GS; 26.iii.2010, 4 ♂♂ 2 ♀♀, CR. Noualhieria pieltaini Gomez-Menor, 1924 (Fig. 1) Bco. Herradura, 12.vi.2011, 1 ♂, GS. Montaña Raijada, 24.iii.2010, 1 ♂ 2 ♀♀, on Sonchus, CR. Puntagorda, 25.iii.2010, 22 ♂♂ 13 ♀♀, from leaf crowns of Sonchus, CR, GS. San Nicolás, 24.iii.2010, 19 ♂♂ 12 ♀♀, on Sonchus (Fig. 1), CR, GS.

Plinthisus (Plinthisus) canariensis Wagner, 1963 El Pinar, 25.iii.2009, 6 ♂♂ 8 ♀♀, BA. Beosus maritimus (Scopoli, 1763) Bco. del Jorado, 23.iii.2009, 2 ♂♂, BA. Bco. de la Galga, 1.v.2009, 1 ♀, GS. El Palmar, 27.iii.2009, 1 ♀, BA. El Paso, farmland, 7.v.2009, 2 ♂♂2 ♀♀, CR. El Paso, gravel pit, 27.iii.2010, 1 ♀, GS. Fuencaliente N, 4.v.2009, 1 ♂, GS. Nogales, 6–7.iii.2008, 1 ♂, JD (ZMAN). Puerto Naos, 24.iii.2010, 1 ♂, GS, 29.iii.2010, 1 ♂, GS. Peritrechus gracilicornis Puton, 1877 La Lomada, 18.vi.2007, 1 ♀, HG. Laguna de Barlovento, 1.iv.2009, 2 ♂♂ 2 ♀♀, BA; 21.iv.2012, 2 ♂♂ 2 ♀♀, HG. San Isidro, 6.v.2009, 2 ♀♀, CR. Raglius alboacuminatus alboacuminatus (Goeze, 1778) Bco. Hondo, 30.iv.2009 1 ♂ 1 ♀, CR; 2.v.2009, 1 ♂ 2 ♀♀, HG. Laguna de Barlovento, 1.iv.2009, 2 ♂♂ 4 ♀♀, BA. Comment. All specimens belong to var. funereus Puton, 1878.

Xanthochilus saturnius (Rossi, 1790) El Paso, farmland, 11.vi.2007, 2 ♂♂ 1 ♀, HG, GS; 7.v.2009, 1 ♂, HG; 11.v.2009, 4 ♂♂ 5 ♀♀, HG, CR. Las Tricias, 19.iv.2012, 1 ♂, HG. Puntagorda, 25.iii.2010, 3 ♀♀, CR, from leaf crowns of Sonchus. Hyalochilus ovatulus (A. Costa, 1853) Bco. Herradura, 12.vi.2011, 1 ♂, GS. Jedey, 11.vi.2011, 1 ♂, GS. La Lomadita, 21.iii.2010, 2 ♂♂ 1 ♀, CR. Los Llanos de Aridane, 26.iii.2009, 1 ♀, BA. Stygnocoris barbieri Péricart, 1993 Fuencaliente N, 7.v.2009, 1 ♀, CR. San Isidro, 6.v.2009, 9 ♂♂ 1 ♀, HG. Volcán Martín, 10.v.2009, 1 ♀, HG.

BERYTIDAE Neides aduncus Fieber, 1859 Bco. Hondo, 30.iv.2009, 1 ♂, CR. El Paso, farmland, 24.iv.2012, 1 ♀, HG. Montaña las Moraditas, 21.iii.2009, 3 ♂♂, BA.

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Berytinus (Berytinus) hirticornis pilipes (Puton, 1875) Bco. de la Galga, 1.v.2009, 1 ♂, CR; 18.iv.2012, 1 ♂, HG. Fuencaliente, 26.ii.2001, 1 ♂, HG. Laguna de Barlovento, 21.iv.2012, 1 ♂, HG. Lomo Oscuro, 9.v.2009, 1 ♂, HG. Montaña Azufre, 27.iii.2010, 1 ♂ 1 ♀, GS. San Simón, 4.v.2009, 1 ♂, CR; 27.iii.2010, 3 ♂♂ 3 ♀♀, CR.

Berytinus (Lizinus) montivagus (Meyer-Dür, 1841) Bco. Hondo, 30.iv.2009, 3 ♂♂ 2 ♀♀, CR, GS; 2.v.2009, 5 ♂♂ 7 ♀♀, HG, CR; 20.iii.2010, 1 ♀, CR. El Paso, farmland, 11.vi.2007, 1 ♂, HG; 7.v.2009, 3 ♂♂ 2 ♀♀, HG, CR; 11.v.2009, 13 ♂♂ 14 ♀♀, HG, CR, GS; 22.iii.2010, 1 ♂, CR. El Pilar, 17.vi.2007, 1 ♂ 2 ♀♀, HG. Las Tricias, 8.v.2009, 1 ♂, HG. Los Llanos, 20.iii.2005, 2 ♀♀, HG. Montaña Azufre, 27.iii.2010, 1 ♂, CR. Montaña las Moraditas, 21.iii.2009, 3 ♀♀. San Simón, 27.iii.2010, 4 ♂♂ 6 ♀♀, BA, CR. Volcán Martín, 10.v.2009, 2 ♀♀, HG.

PYRRHOCORIDAE Scantius aegyptius aegyptius (Linnaeus, 1758) La Lomadita, 21.iii.2010, 1 ♂ 1 ♀, GS. Puerto Naos, 24.iii.2010, 1 ♂ 3 ♀♀, GS; 29.iii.2010, 1 ♂, GS.

STENOCEPHALIDAE agilis agilis (Scopoli, 1763) Lomo Oscuro, 9.v.2009, 1 ♀, HG. Los Llanos de Aridane, swimming pool, 22.iii.2009, 4 ♂♂, BA. San Simón, 4.v.2009, 1 ♀, GS.

RHOPALIDAE Brachycarenus tigrinus (Schilling, 1829) El Paso, farmland, 7.v.2009, 3 ♂♂, CR, GS. El Paso, gravel pit, 22.iii.2010, 1 ♂, GS; 27.iii.2010, 2 ♂♂ 1 ♀, GS. Las Tricias, 2.v.2009, 1 ♂, GS. Los Llanos de Aridane, 23.iii.2009, 1 ♂, 29.iii.2009, 1 ♀, BA.

Corizus hyoscyami nigridorsum (Puton, 1874) Bco. del Jorado, 23.iii.2009, 1 ♂, BA; 13.vi.2011, 1 ♂, GS. Bco. de las Angustias, 9.iii.2005, 1 ♂, HG. Bco. Hondo, 30.iv.2009, 4 ♂♂ 5 ♀♀, on Sida rhombifolia, CR; 2.v.2009, 1 ♂ 3 ♀♀, HG. El Paso, farmland, 11.v.2009, 1 ♂ 1 ♀, HG, CR; 22.iii.2010, 2 ♂♂ 2 ♀♀, CR, GS. El Paso, gravel pit, 23.iii.2010, 1 ♀, CR; 25.iii.2010, 2 ♂♂ 1 ♀, CR; 27.iii.2010, 3 ♂♂ 2 ♀♀, GS; 5.vi.2011, 1 ♀, GS. El Pilar, 2.v.2009, 1 ♂, GS. La Punta, 2.iv.2009, 1 ♂, BA. Las Tricias, 7.v.2011, 1 ♂, GS. Lomo Oscuro, 23.iv.2012, 1 ♂ 2 ♀♀, HG. Montaña la Laguna, 22.iii.2010, 4 ♂♂, CR, GS. Puerto Naos, 29.iii.2010, 1 ♂ 1 ♀, CR. Puntagorda, 10.iii.2008, 2 ♂♂ 2 ♀♀; 17.iii.2008, 2 ♂♂ 2 ♀♀, JD (ZMAN). Puntagorda, Puerto, 25.iii.2010, 1 ♂, CR. Tazacorte, 21.iii.2010, 2 ♂♂, GS. Torre del Time, 1.vi.2011, 7 ♂♂ 2 ♀♀, GS.

Liorhyssus hyalinus (Fabricius, 1794) Bco. Hondo, 2.v.2009, 1 ♂, HG. El Paso, farmland, 11.vi.2007, 1 ♂, HG; 11.v.2009, 2 ♀♀, CR. La Lomadita, 21.iii.2010, 1 ♀ 1 larva 5 → 1 ♀, CR, GS. La Punta, 2.iv.2009, 1 ♂, BA. Las Tricias, 8.v.2009, 1 ♀, GS; 7.v.2011, 2 ♀♀, GS. Lomo Oscuro, 9.vi.2007, 1 ♂, HG; 9.v.2009, 1 ♂, GS. Los Llanos de Aridane, 23.iii.2009, 1 ♀, BA. Puerto Naos, 24.iii.2010, 2 ♂♂ 1 ♀ 5 larvae 5 → 1 ♂, CR, GS; 29.iii.2010, 1 ♀ 1 larva 5 → 1 ♀, CR, GS. Puntagorda, 17.iii.2008, 1 ♂, JD (ZMAN). Tazacorte, 21.iii.2010, 1 ♀, GS.

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ALYDIDAE Camptopus lateralis (Germar, 1817) El Paso, farmland, 7.v.2009, 1 ♂ 3 ♀♀, HG, CR; 11.v.2009, 2 ♂♂ 1 ♀, HG, CR, GS; 22.iii.2010, 1 ♀, GS. El Paso, gravel pit, 11.v.2009, 3 ♂♂ 3 ♀♀; 22.iii.2010, 1 ♀; 23.iii.2010, 1 ♂ 1 ♀, CR; 27.iii.2010, 1 ♂, GS. Fuencaliente N, 20.iii.2010, 1 ♀, CR. Laguna de Barlovento, 21.iv.2012, 2 ♀♀, HG. Las Tricias, 8.v.2009, 1 ♀, HG; 7.v.2011, 1 ♀, GS. Los Llanos de Aridane, 22.iii.2009, 1 ♂, BA. Montaña de la Negra, 25.iii.2010, 1 ♂, CR. Montaña la Laguna, 22.iii.2010, 2 ♂♂, CR, GS. Torre del Time, 1.vi.2011, 1 ♀, GS.

COREIDAE Arenocoris waltlii (Herrich-Schaeffer, 1835) Bco. Hondo, 30.iv.2009, 1 ♂ 2 ♀♀, GS; 2.v.2009, 2 ♂♂ 1 larva → 1 ♂, HG, CR. El Paso, farmland 22.iii.2010, 3 ♂♂ 1 ♀, CR. El Paso, gravel pit, 25.iii.2010, 1 ♂, CR; 27.iii.2010, 1 ♀, GS; 5.vi.2011, 1 ♂ 1 ♀, GS. Jedey, 11.vi.2011, 1 ♂, GS. Los Llanos de Aridane, 22.iii.2009, 2 ♀♀, BA. Montaña las Moraditas, 21.iii.2009, 1 ♀, BA. Santo Domingo, 21.iii.2010, 1 ♀, CR, GS. 27.iii.2010, 1 ♀; 5.vi.2011, 1 ♂ 1 ♀, GS.

Coriomeris affinis (Herrich-Schaeffer, 1839) Bco. de las Angustias, 20.iii.2005, 1 ♂, HG; 3.vi.2011, 1 ♂, GS. Bco. Hondo, 30.iv.2009, 1 ♀, GS; 2.v.2009, 1 ♂ 3 ♀♀, HG, CR, GS. El Palmar, 11.iii.2008, 1 ♂, JD (ZMAN); 27.iii.2009, 1 ♀, BA. El Paso, farmland, 7.v.2009, 3 ♂♂ 1 ♀, CR, GS; 11.v.2009, 1 ♂ 1 ♀, HG, GS; 22.iii.2010, 6 ♂♂ 2 ♀♀, CR, GS. El Paso, gravel pit, 22–23.iii.2010, 5 ♂♂ 1 ♀, CR, GS; 5.vi.2011, 1 ♂ 1 ♀, GS. El Pinar, 25.iii.2009, 3 ♂♂, BA. Montaña las Moraditas, 21.iii.2009, 1 ♂ 1 ♀, BA. La Fajana, 9.iii.2008, 1 ♂, JD (ZMAN). La Punta, 25.iv.2009, 1 ♂. Las Tricias, 7.v.2011, 1 ♀, GS. Lomo Oscuro, 20.iii.2010, 1 ♂, GS. Montaña Azufre, 27.iii.2010, 1 ♀, GS. Montaña de la Breña, 28.iii.2009, 1 ♀, BA. Puerto Naos, 24.iii.2010, 1 ♂ 1 ♀, GS. Puntagorda, 17.iii.2008, 6 ♂♂ 1 ♀, JD (ZMAN). San Isidro, 6.v.2009, 9 ♂♂ 1 ♀, CR, GS. Santo Domingo, 21.iii.2010, 1 ♂, CR; 26.iii.2010, 1 ♀, CR. Tijarafe, 16.iii.2008, 1 ♂, JD (ZMAN). Torre del Time, 1.vi.2011, 1 ♀, GS.

Haploprocta sulcicornis (Fabricius, 1794) Bco. del Jorado, 23.iii.2009, 1 ♀, BA. Bco. de las Angustias, 5.v.2009, 1 ♀, GS. Bco. Hondo, 2.v.2009, 1 ♂, HG. El Paso, farmland, 11.v.2009, 1 ♂, GS. Fuencaliente, 21.iii.2005, 1 ♂ 1 ♀, HG. Fuencaliente N, 4.v.2009, 1 ♂, CR, under Rumex lunaria, 7.v.2009, 1 ♂, GS. Lomo Oscuro, 20.iii.2010, 1 ♀, CR. Puntagorda, 23.iii.2010, 1 ♀, GS; 17.iii.2008, 1 ♀, JD (ZMAN). Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS. Volcán Martín, 10.v.2009, 1 ♀, HG.

Syromastus rhombeus (Linnaeus, 1767) Bco. de la Galga, 1.v.2009, 1 ♀, GS; 28.iii.2010, 1 ♂ 1 ♀, CR. Bco. Hondo, 30.iv.2009, 1 ♂, GS; 2.v.2009, 1 ♂, HG. El Paso, farmland, 7.v.2009, 1 ♂ 1 ♀, CR; 22.iii.2010, 2 ♂♂ 1 ♀, CR, GS. El Paso, gravel pit, 23.iii.2010, 1 ♂ 1 ♀, CR, GS. Hoya Grande, 8.iii.2008, 1 ♀, JD (ZMAN). Lomo Oscuro, 20.iii.2010, 1 ♂, CR, 7.vi.2011, 1 ♂, GS. Montaña las Moraditas, 21.iii.2009, 1 ♂, BA. Puntagorda, 17.iii.2008, 1 ♂ 1 ♀, JD (ZMAN). Volcán Martín, 10.v.2009, 1 ♀, HG.

CYDNIDAE Cydnus aterrimus (Forster, 1771) Montaña Argual, 29.iii.2009, 1 ♂, BA. Montaña de la Breña, 28.iii.2009, 1 ♀, BA. La Lomadita, 21.iii.2010, 1 ♀, GS. San Simón, 4.v.2009, 1 ♂ 2 ♀♀, CR, under Euphorbia.

476 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Heteroptera of La Palma, Canary Islands

Macroscytus brunneus (Fabricius, 1803) La Fajana, 9.iii.2008, 1 ♂, JD (ZMAN). Lomo Oscuro, 9.v.2009, 1 ♀, HG; 23.iv.2012, 1 ♂, HG. Los Llanos de Aridane, 22.iii.2009, 1 ♂, in swimming pool, BA. Santo Domingo, 26.iii.2010, 2 ♀♀, CR.

Crocistethus aeneus (Brullé, 1839) El Pinar, 25.iii.2009, 1 ♂, BA.

SCUTELLERIDAE Odontotarsus caudatus (Burmeister, 1835) El Paso, farmland, 22.iii.2010, 1 ♂ 1 ♀, CR. El Paso, gravel pit, 23.iii.2010, 7 ♂♂ 5 ♀♀, CR, GS; 25.iii.2010, 2 ♂♂ 2 ♀♀, CR; 27.iii.2010, 1 ♂ 1 ♀, GS. Puntagorda, 17.iii.2008, 1 ♂ 1 ♀, JD (ZMAN).

Odontoscelis (Odontoscelis) dorsalis (Fabricius, 1798) Bco. Hondo, 30.iv.2009, 1 ♀, GS; 2.v.2009, 2 ♂♂, HG. El Paso, farmland, 11.vi.2007, 1 ♂ 1 ♀, HG; 7.v.2009, 1 ♂, CR; 11.v.2009, 9 ♂♂ 2 ♀♀, HG, CR, GS; 22.iii.2010, 5 ♂♂ 2 ♀♀, CR, GS. El Paso, gravel pit, 25.iii.2010, 1 larva, CR. La Punta, 2.iv.2009, 1 ♀, BA. Puntagorda, 17.iii.2008, 1 larva 5, JD (ZMAN). San Simón, 27.iii.2010, 1 ♂, CR. Santo Domingo, 21.iii.2010, 1 ♂, CR; 26.iii.2010, 1 ♂, CR.

PENTATOMIDAE Dolycoris numidicus Horváth, 1907 El Paso, gravel pit, 5.vi.2011, 1 ♀, GS. Las Tricias, 8.v.2009, 1 ♀, HG; 7.v.2011, 1 ♀, GS. Puntagorda, 17.iii.2008, 1 ♀, JD (ZMAN).

Eudolycoris alluaudi (Noualhier, 1893) Bco. Hondo, 30.iv.2009, 1 ♂, HG. Fuencaliente N, 4.v.2009, 6 ♂♂ 18 ♀♀, CR, GS; 7.v.2009, 3 ♂♂ 9 ♀♀, CR, GS; 10.v.2009, 2 ♂♂ 3 ♀♀, on Sideritis canariensis, HG, CR; Jedey, 11.vi.2011, 2 ♀♀, GS; Roque de los Muchachos, 20.iv.2012, 1 ♂, HG; Volcán San Martín, 10.v.2009, 2 ♂♂ 5 ♀♀, HG.

Eysarcoris ventralis (Westwood, 1837) Bco. de la Galga, 9.vi.2007, 1 ♂ 2 ♀♀, HG; 1.v.2009, 4 ♂♂ 5 ♀♀, HG, CR, GS. Bco. de las Angustias, 16.iv.2012, 1 ♀, HG. Laguna de Barlovento, 21.iv.2012, 1 ♂ 1 ♀, HG. Puerto de Tazacorte, Bco. de las Angustias, 14.iii.2008, 2 ♀♀, JD (ZMAN). Puerto Naos, 24.iii.2010, 5 larvae → 1 ♂ 3 ♀♀, CR; 29.iii.2010, 1 ♀, GS.

Holcogaster fibulata (Germar, 1831) El Pilar, 3.v.2009, 2 ♂♂ 1 ♀, HG, CR; 23.iii.2010, 15 ♂♂ 12 ♀♀, on Pinus canariensis, CR, GS. Montaña las Moraditas, 21.iii.2009, 1 ♀, on Pinus canariensis, BA. Pico Birigoyo, 9.vi.2011, 1 ♂, GS.

Mecidea lindbergi Wagner, 1954 Puerto de Tazacorte, 13.iii.2008, 6 ♂♂ 5 ♀♀, JD (ZMAN). Puntagorda, 10.iii.2008, 8 ♂♂ 6 ♀♀, JD (ZMAN).

Mecidea pallidissima Jensen-Haarup, 1922 Puerto de Tazacorte, 13.iii.2008, 1 ♂, JD (ZMAN). San Simón, 27.iii.2010, 1 ♀, GS.

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Acrosternum rubescens (Noualhier, 1893) Los Llanos de Aridane, 20.iii.2009, 4 ♂♂ 8 ♀♀, on Euphorbia, BA. Nogales, 6–7.iii.2008, 1 ♂ 1 ♀, JD (ZMAN).

Nezara viridula (Linnaeus, 1758) El Paso, gravel pit, 27.iii.2010, 2 ♂♂, GS. La Lomadita, 21.iii.2010, 1 ♂ 1 ♀, CR, GS. Las Tricias, 7.v.2009, 1 ♀, CR. Lomo Oscuro, 7.vi.2011, 1 ♀, GS. Montaña Azufre, 27.iii.2010, 1 ♀, GS. Puerto Naos, 24.iii.2010, 2 ♂♂ 1 ♀, CR, GS; 26.iii.2010, 1 ♂ 1 ♀, CR.

Piezodorus punctipes Puton, 1889 Barlovento, 21.iii.2010, 2 ♀♀, GS. Bco. de las Angustias, 5.v.2009, 1 ♀, GS. Bco. Hondo, 30.iv.2009, 1 ♂, on Adenocarpus; 2.v.2009, 1 ♂, CR. El Paso, farmland, 7.v.2009, 8 ♂♂ 1 ♀, GS; 11.v.2009, 1 ♂ 2 ♀♀, CR, GS. El Paso, gravel pit, 22.iii.2010, 1 ♀, GS. El Pilar, 3.v.2009, 9 ♂♂ 30 ♀♀, CR, GS; 23.iii.2010, 10 ♂♂ 6 ♀♀, on Adenocarpus, CR, GS. Fuencaliente N, 10.v.2009, 1 ♀. Pico Birigoyo, 9.vi.2011, 1 ♀, GS. Torre del Time, 1.vi.2011, 1 ♀, GS.

Sciocoris (Neosciocoris) sideritidis Wollaston, 1858 Bco. del Jorado, 10.vi.2011, 1 ♀, GS; 13.vi.2011, 1 ♂, GS. Bco de las Angustias, 20.iii.2005, 1 ♂, HG. Bco. Herradura, 12.vi.2011, 1 ♂, GS. Bco. Hondo, 30.iv.2009, 10 ♂♂ 2 ♀♀, HG, CR; 2.v.2009, 5 ♂♂ 4 ♀♀, HG, CR; 20.iii.2010, 1 ♂, CR. El Palmar, 27.iii.2009, 2 ♀♀, BA. El Paso, farmland, 11.vi.2007, 1 ♀, HG. Fuencaliente, 9.vi.2007, 1 ♀, HG. Fuencaliente N, 7.v.2009, 53 ♂♂ 21 ♀♀, CR, GS; 10.v.2009, 2 ♀♀, on Sideritis canariensis, CR. Jedey, 11.vi.2011, 3 ♂♂ 7 ♀♀, GS. Mazo, 8.iv.2000, 1 ♀, H. Ziegler leg. (GS). Nogales, 6–7.iii.2008, 1 ♀, JD (ZMAN). San Nicolás, 24.iii.2010, 3 ♂♂ 3 ♀♀, CR, GS. San Simón, 27.iii.2010, 1 ♀, CR. Torre del Time, 1.vi.2011, 1 ♀, GS. Volcán Martín, 21.ii.2001, 1 ♂, HG; 10.v.2009, 1 ♂, HG.

Sciocoris (Parasciocoris) canariensis Lindberg, 1953 El Paso, farmland, 2.v.2009, 5 ♂♂ 5 ♀♀, GS. La Fajana, 9.iii.2008, 2 ♂♂, JD (ZMAN). Lomo Oscuro, 9.v.2009, 1 ♂, HG. Los Llanos de Aridane, 20.iii.2009, 6 ♂♂ 4 ♀♀; 26.iii.2009, 1 ♂, on Forskalea, BA. Mazo, 5.iii.1994, 1 ♂ 1 ♀, P. Sprick leg. (HG); Montaña la Laguna, 13.vi.2007, 1 ♀, HG; 2.v.2009, 18 ♂♂ 7 ♀♀, HG, CR, GS, under Forskalea. Nogales, 6–7.iii.2008, 1 ♂, JD (ZMAN). Puerto Naos, 24.iii.2010, 6 ♂♂ 6 ♀♀, GS; 26.iii.2010, 4 ♂♂ 5 ♀♀ 1 larva, under Forskalea, CR. Puntagorda, Puerto, 25.iii.2010, 2 ♀♀, CR. San Nicolás, 24.iii.2010, 1 ♂, CR. Tazacorte, 21.iii.2010, 1 ♂ 2 ♀♀, GS.

Sciocoris (Sciocoris) angularis Puton, 1889 Bco. de las Angustias, 20.iii.2005, 1 ♂ 1 ♀, HG. Bco. Hondo, 30.iv.2009, 2 ♂♂ 3 ♀♀, GS; 2.v.2009, 1 ♂ 2 ♀♀, HG, GS. El Paso, farmland, 7.v.2009, 4 ♂♂ 8 ♀♀, HG, CR, GS; 11.v.2009, 2 ♀♀, CR; 22.iii.2010, 1 ♀, GS. La Fajana, 9.iii.2008, 1 ♀, JD (ZMAN). Las Tricias, 14.iii.2005, 2 ♂♂ 2 ♀♀, HG. Los Llanos de Aridane, 20.iii.2009, 3 ♂♂ 1 ♀; 26.iii.2009, 1 ♀, BA. Mazo, 1.vii.1989, 1 ♂, E. Heiss leg. (GS). Pico de la Nieve, 1890 m a.s.l., 6.v.2009, 1 ♂ 2 ♀♀, HG. Puntagorda, 15.iii.2008, 1 ♀; 17.iii.2008, 1 ♂ 1 ♀, JD (ZMAN). Salinas, 12.vi.2007, 1 ♀. Torre del Time, 1.vi.2011, 1 ♂ 1 ♀, GS. Volcán Martín, 1165 m a.s.l., 10.v.2009, 1 ♀, CR in ash/fine sand.

Sciocoris (Sciocoris) helferii Fieber, 1851 Nogales, 6–7.iii.2008, 1 ♂, JD (ZMAN). Puntagorda, 17.iii.2008, 2 ♀♀, JD (ZMAN). Santo Domingo, 2.2 km SW, 26.iii.2010, 2 ♂♂ 1 ♀, CR.

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Figs 1–3. 1 – Sonchus sp., San Nicolás, potential food plant of Noualhieria pieltaini Gómez-Menor, 1924. 2 – Rumex lunaria near Torre del Time (1.vi.2011): food plant of Haploprocta incognita Heiss et Moulet, 1994 and Lindbergopsallus lunariae Rieger, Strauss et Günther, 2011. 3 – Haploprocta incognita Heiss et Moulet, 1994, Torre del Time (1.vi.2011).

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Eurydema (Eurydema) lundbladi Lindberg, 1960 El Pinar, 25.iii.2009, 1 ♂, BA. Las Tricias, 7.v.2011, 1 ♀, GS. Pico de la Nieve, 1890 m a.s.l., 6.v.2009, 59 ♂♂ 58 ♀♀, on Descurainia, HG, CR, GS. Roque de los Muchachos, 27.ii.2001, 6 ♂♂ 3 ♀♀, HG; 20.iv.2012, 3 ♂♂ 1 ♀, HG.

Eurydema (Eurydema) ornata (Linnaeus, 1758) El Pinar, 25.iii.2009, 1 ♀, BA. El Paso, farmland, 7.v.2009, 6 ♂♂ 4 ♀♀, CR, GS; 11.v.2009, 3 ♂♂ 1 ♀, HG, CR, GS. El Paso, gravel pit, 22.iii.2010, 1 ♀, GS; 27.vii.2010, 1 ♂ 1 ♀, GS. Las Tricias, 8.v.2009, 2 ♀♀, CR; 7.v.2011, 1 ♂, GS. Puntagorda, 17.iii.2008, 1 ♀, JD (ZMAN).

Ancyrosoma leucogrammes (Gmelin, 1790) San Simón, 4.v.2009, 1 ♂, CR.

Graphosoma (Graphosoma) interruptum A. White, 1839 Bco. del Jorado, 13.vi.2011, 4 ♂♂, GS. Bco. Herradura, 12.vi.2011, 3 ♀♀, GS. El Paso, gravel pit, 5.vi.2011, 1 ♂, GS. Torre del Time, 1.vi.2011, 1 ♂, GS.

Concluding remarks In this contribution 28 species are listed for the first time from La Palma (! in Table 2), of which six are new to the Canary Islands as well (* in Table 2). For eight species the occurrence on La Palma is confirmed ( in Table 2). With these additions the total number of known species from the Canary Islands is raised to 400 and 216 species are now listed from La Palma. Of the species known from La Palma, 48 (22.2 %) are Canarian endemics, and four of these are only known from La Palma: Lindbergopsallus lauri and L. lunariae (Miridae), Collartida tanausu (Reduviidae) and Nysius gloriae (Lygaeidae).

Acknowledgements We would like to thank Dr. E. Heiss (Innsbruck, Austria), Dr. F. Lange (Niedernhausen, Germany), Mr. W. Löderbusch (Markdorf, Germany), Dr. P. Sprick (Hannover, Germany) and Dr. H. Ziegler † (Biberach, Germany) for the transfer of data and material.

References

AUKEMA B. 2012: New records of Coranus kiritshenkoi Bergevin, 1932 from the Canary Islands (Heteroptera: Reduviidae). Entomologische Zeitschrift (Stuttgart) 122(3): 135–136. AUKEMA B., DUFFELS J. P. & BÁEZ M. 2006: A checklist of the Heteroptera of the Canary Islands (Insecta). Pp. 755–774. In: RABITSCH W. (ed.): Hug the bug. For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss. Denisia 19:.1–1184. AUKEMA B., RIEGER CH. & RABITSCH W. 2013: Catalogue of the Heteroptera of the Palaearctic Region 6. Supplement. Netherlands Entomological Society, Amsterdam, xxiii + 629 pp.

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BÁEZ M. & ZURITA N. 2001: Hemiptera Heteroptera. Pp. 182–197. In: IZQUIERDO I., MARTÍN J. L., ZURITA N. & ARECHAVALETA M. (eds): Lista de especies silvestres de Canarias (hongos, plantas y animales terrestres) 2001. Consejería de Medio Ambiente y Ordenación Territorial, Gobierno de Canarias, 437 pp. BÁEZ M., MARTÍN E. & ZURITA N. 2004: Hemiptera Heteroptera. Pp. 192–207. In: IZQUIERDO I., MARTÍN J. L., ZURITA N. & ARECHAVALETA M. (eds): Lista de especies silvestres de Canarias (hongos, plantas y animales terrestres) 2004. Consejería de Medio Ambiente y Ordenación Territorial, Gobierno de Canarias, 500 pp. HEISS E. 1997: Nachtrag zur Heteropterenfauna der Kanarischen Inseln V (Insecta, Heteroptera). Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 84: 359–369. FARACI F. 2012: Presencia de Sigara (Halicorixa) selecta (Fieber, 1848) en Fuerteventura (Islas Canarias) (Hemiptera: Heteroptera: Corixidae). Heteropterus Revista de Entomología 11: 241–243 LINDBERG H. 1953: Hemiptera Insularum Canariensium. Systematik, Ökologie und Verbreitung der Kanarischen Heteropteren und Cicadinen. Commentationes Biologicae 14(1): 1–304. LINDBERG H. 1960: Supplementum Hemipterorum Insularum Canariensium. Commentationes Biologicae 22(6): 1–20. LIS B. 2003: Revision of the genus Cantacader Amyot et Serville, 1843 (Hemiptera: Heteroptera: Cantacaderidae). Polskie Pismo Entomologiczne 72(Suppl.): 1–222. NOUALHIER M. 1893: Voyage de M. Ch. Alluaud aux îles Canaries (Novembre 1889 – Juin 1890). 2e Mémoire. Hémiptères Gymnocérates & Hydrocorises. Annales de la Société Entomologique de France 62: 5–18. PUTSHKOV P. V. & M OULET P. 2010: Hémiptères Reduviidae d’Europe occidentale. Faune de France 92: 1–668. RIBES J. & PAGOLA-CARTE S. 2008: Kalama montisclari n. sp.: Un nuevo tíngido de la Islas Canarias (Hemiptera: Heteroptera: Tingidae). Heteropterus Revista de Entomología 8: 131–135. RIBES J., PAGOLA-CARTE S. & HEISS E. 2008: Two new Phylinae (Hemiptera: Heteroptera: Miridae) from the Canary Islands. Acta Entomologica Musei Nationalis Pragae 48: 423–431. RIEGER CH. 2012: Eine weitere Art von Lindbergopsallus Wagner, 1962, von der Kanareninsel La Palma (Heteroptera: Miridae). Entomologische Zeitschrift (Stuttgart) 122: 273–275. RIEGER CH., STRAUß G. & GÜNTHER H. 2011: Ein neuer Lindbergopsallus Wagner, 1961 von der Kanareninsel La Palma (Heteroptera: Miridae). Entomologische Zeitschrift (Stuttgart) 121: 255–258. SANTAMARÍA Á., GUTIÉRREZ J., RÉGIL J. A. & PÉREZ DELGADO A. J. 2012: Nuevos registros referentes a la distribución de los hemípteros acuáticos (Nepomorpha y Gerromorpha) de las islas Canarias (España). Boletín de la Sociedad Entomológica Aragonesa 50: 555–558.

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Table 1. List of collecting sites on La Palma. To avoid constant repetition of longer names of collecting sites, the shortened forms listed below in bold type are used in the text. (Continued on pages 483–485.)

Barlovento Laurisilva 2.8 km SW of Barlovento (770 m) 17°49′27″ W 28°48′36″ N

Bco. de la Galga Bco. de la Galga between visitor’s center and Cubo de la Galga (340–780 m) Cubo 17°46′41″ W 28°45′29″ N Visitor’s center 17°46′11″ W 28°45′01″ N

Bco. del Agua 17°48′19″ W 28°47′11″ N SW of Los Tilos (530 m)

Bco. de las Angustias 17°52′ W 28°42′ N Los Llanos, Bco. de las Angustias between parking place and Dos Aguas (580 m)

Bco. Gallegos Bco. between Barlovento and Gallegos (800 m) 17°49′59″ W 28°48′22″ N

Bco. Herradura 17°46′33″ W 28°48′41″ N Bco. N of Los Sauces (600 m)

Bco. Hondo Bco. between Fuencaliente and Las Manchas (730–740 m) 17°51′59″ W 28°32′59″ N

Bco. de Jorado 17°57′08″ W 28°42′19″ N Bco. S of Tijarafe (500–700 m)

Breña Alta 17°48′40″ W 28°39′24″ N Surrounding of Túnel Chico (800 m)

Callejones Outskirts SE of Callejones (290 m) 17°46′06″ W 28°36′13″ N

El Pinar 17°55′ W 28°41′ N Between Cruz del Llano (1020 m) and Torre del Time (1180 m)

El Palmar 17°56′ W 28°50′ N Between Santo Domingo de Garafia and El Palmar (200–360 m)

El Paso, gravel pit 17°50′44″ W 28°39′42″ N Gravel pit NE of El Paso near Ermita Virgen del Pino (820–850 m)

El Paso, farmland 17°50′13″ W 28°38′50″ N Pasture, fields and fallow land E of El Paso (940 m)

El Pilar Cumbre Vieja between refugio El Pilar and refugio at the foot of Montaña Andrés Martín (1440–1510 m) El Pilar 17°50′10″ W 28°36′34″ N Refugio 17°49′19″ W 28°35′26″ N

El Tablado 17°53″ W 28°48′ N W of Roque Faro (1050 m)

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Faro de Barlovento 17°46′42″ W 28°50′18″ N Ruderal vegetation near the lighthouse (30 m)

Fuencaliente N 17°51′46″ W 28°31′17″ N Tracks along the fields W of the road Fuencaliente Las Manchas (750 m)

Fuencaliente NE 17°49′17″ W 28°31′06″ N Waste site at the roadside NE of Fuencaliente (770 m)

Fuencaliente ca 17°52′ E 28°33′ E Cultivated/derelict farmland between Fuencaliente and Las Manchas (740 m)

Hoya Grande (El Pinar) (1490 m) 17°54′58″ W 28°42′17″ N

Jedey 17°52′38″ W 28°34′19″ N Old lava flow with sparse vegetation, approximately 1.5 km S of Jedey

Laguna de Barlovento 17°48′27″ W 28°48′28″ N Surrounding of the water reservoir S of Barlovento (760 m)

La Fajana 17°51′ W 28°50′ N Near Franceses (400 m)

La Lomadita 17°46′28″ W 28°49′50″ N Ruderal site 0.8 km NE of La Lomadita near Punta Cumplida (30 m)

La Punta 17°56′ W 28°39′ N From Mirador del Time towards Puerto Tazacorte

Las Lomadas ca 17°46′ W 28°47′ N Laurisilva pr. Las Lomadas (500 m)

Las Tricias 17°58′28″ W 28°47′34″ N Farmland SW of Las Tricias (dragon trees) (470–770 m)

Lomo Oscuro 17°46′07–08″ W Derelict, partly terraced farmland 1 km SE of Lomo Oscuro 28°34′29–34″ N respectively 1.5 km NE of Malpaíses (150 m)

Los Barros 17°54′ W 28°41′ N From Lomo de Los Caballos into Bco. de las Angustias

Los Llanos de Aridane Farmland at the northern outscirts of Los Llanos (450 m) 17°54′58″ W 28°40′19″ N Water basin 17°54′15″ W 28°39′42″ N

Mirca 17°46′05″ W 28°42′21″ N Derelict farmland at the northern outskirts of Mirca (200m)

Montaña Argual (320 m) 17°55′22″ W 28°39′21″ N

Montaña Azufre 17°46′27″ W 28°33′49″ N Foot of the Volcano 1 km SE of Malpaíses, derelict farmland (190 m)

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Montaña de la Breña 17°46′ W 28°37′ N Foothpath to Villa de Mazo (560 m)

Montaña de la Negra 17°59′17″ W 28°47′03″ N Puntagorda, foot of the Montaña Negra, goats grazing (460 m)

Montaña la Laguna 17°54′51″ W 28°37′44″ N Volcano W of La Laguna (275–315 m)

Montaña las Moraditas 17°51′24″ W28′38′50″ N Pasture (900 m)

Montaña Raijada 17°52′35″ W 28°36′42″ N Vulcano N of San Nicolas (690 m)

Monte de Luna (735 m) 17°48′57″ W 28°32′05″ N

Nogales 17°44′ W 28°45′ N NE of San Juan de Puntallana (100 m)

Pico Birigoyo 17°50′19″ W 28°36′29″ N Firebreak on the northeastern mountainside of P. Birigoyo (1500–1600 m)

Pico de la Nieve Eastern slope 1890 m (pine forest) 17°49′31″ W 28°44′06″ N 1070 m (pine forest) 17°47′29″ W 28°43′10″ N 665 m (laurisilva) 17°46′21″ W 28°43′17″ N

Puerta de Tazacorte 17°56′ W 28°39′ N Bco. de las Angustias (25 m)

Puerto Naos 17°54′27″ W 28°35′18″ N Outskirts of Puerto Naos, ruderal site/garbage dump (20 m)

Punta Bacha de San Simón 17°45′41″ W 28°34′32″ N Grove of Tamarix surrounded by derelict farmland (10 m)

Puntagorda 17°59′23″ W 28°46′09″ N Farmland at the eastern outskirts of Puntagorda (580 m)

Puntagorda, Puerto 17°58′10″ W 28°49′02″ N Derelict farmland near Puerto (130–160 m)

Roque de los Muchachos (2300 m) 17°52′52″ W 28°45′42″ N

Salinas 17°50′28″ W 28°27′15″ N Saltworks S of Fuencaliente (10–20 m)

San Isidro 17°48′37″ W 28°37′51″ N Farmland/Woodland S of San Isidro (700 m)

San Nicolás 17°52′38″ W 28°36′26″ N Slope NE of San Nicolás, derelict farmland (660 m)

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San Simón 17°45′40″ W 28°35′31″ N Pasture 0.7 km SSW of San Simón (150 m)

Santa Cruz (30 m) 17°45′55″ W 28°41′00″ N

Santo Domingo 17°57′44″ W 28°48′53″ N Derelict farmland 2.2 km SW of Santo Domingo (325 m)

Tazacorte 17°56′27″ W 28°39′17″ N Surroundings of Puerto Tazacorte (30 m)

Tigalate (700 m) 17°48′07″W 28°33′14″ N

Tijarafe 17°57′14″ W 28°42′25″ N Cemeterio (600 m)

Torre el Time (El Pinar) 17°55′25–28″ W 28°42′15″ N Surroundings of Torre el Time (1150 m)

Villa de Mazo (450 m) 17°46′49″ W 28°37′13″ N

Volcán San Antonio 17°51′ W 28°29′ N Foothpath from Los Quemados (560–620 m)

Volcán San Martín 17°50′36″ W 28°32′12″ N Volcano N of Fuencaliente (1550 m)

Volcán Teneguía 17°50′51″ W 28°28′01″ N Foot of the volcano S of Fuencaliente (240 m)

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Table 2. The Heteroptera of the Canary Islands. Abbreviations used: H – El Hierro, P – La Palma, G – La Gomera, T – Tenerife, C – Gran Canaria, F – Fuerteventura, L – Lanzarote. * – additional species; E – endemic species; – published records (AUKEMA et al., 2006, and additional papers); !– unspecified island records from BÁEZ & ZURITA (2001) and BÁEZ et al. (2004); – confirmed records; ! – new records. (Continued on pages 487–493.)

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Note: In the list of the heteropteran species of the Canary Islands in AUKEMA et al. (2006) Prostemma guttula (Nabidae) and Dicranocephalus agilis (Stenocephalidae) were erroneously reported for Fuerteventura and are removed here from the list for this island. The records were made on Tenerife (Prostemma) and Lanzarote (Dicranocephalus), respectively.

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ISSN 1211-8788 Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 495–541, 2013

New and interesting records of true bugs (Hemiptera: Heteroptera) from the Czech Republic and Slovakia V

PETR KMENT1, KAREL HRADIL2, PETR BAÒAØ3, ONDØEJ BALVÍN4, JOZEF CUNEV5, TOMÁŠ DITRICH6, ZDENÌK JINDRA7, MAGDALENA ROHÁÈOVÁ8, MICHAL STRAKA9 & JAN SYCHRA9 1 Department of Entomology, National Museum, Kunratice 1, CZ-148 00 Praha 4, Czech Republic; e-mail: [email protected] 2 State Phytosanitary Administration, Havlíèkova 56, 506 01 Jièín; e-mail: [email protected] 3 Moravian Museum, Department of Entomology, Hviezdoslavova 29a, CZ-627 00 Brno, Czech Republic; e-mail: [email protected] 4 Department of Zoology, Faculty of Science, Charles University, Vinièná 7, CZ-128 44, Praha 2, Czech Republic; e-mail: [email protected] 5 Piešťanská 14, 949 01 Nitra; e-mail: [email protected] 6 Department of Biology, Faculty of Education, University of South Bohemia, Jeronýmova 10, CZ-371 15 Èeské Budìjovice, Czech Republic; e-mail: [email protected] 7 Department of Plant Protection, Faculty of Agrobiology, Food and Natural Resources, Czech University of Agriculture, CZ-165 21 Praha 6-Suchdol, Czech Republic; e-mail: [email protected] 8 Beskydy Museum, Hluboká 66, CZ-738 01 Frýdek-Místek, Czech Republic; e-mail: [email protected] 9 Department of Botany and Zoology, Faculty of Science, Masaryk University, Kotláøská 2, CZ-611 37 Brno, Czech Republic; e-mails: [email protected], [email protected]

KMENT P. et al. 2013: New and interesting records of true bugs (Hemiptera: Heteroptera) from the Czech Republic and Slovakia V. In: KMENT P., MALENOVSKÝ I. & KOLIBÁÈ J. (eds.): Studies in Hemiptera in honour of Pavel Lauterer and Jaroslav L. Stehlík. Acta Musei Moraviae, Scientiae biologicae (Brno) 98(2): 495–541. – The following species are recorded for the first time from the Czech Republic: Microvelia pygmaea (Dufour, 1833) (from Moravia), Isometopus mirificus Mulsant et Rey, 1879 (Moravia), Tupiocoris rhododendri (Dolling, 1972) (Bohemia), Tuponia macedonica Wagner, 1957 (Moravia), Geocoris erythrocephalus (Lepeletier et Serville, 1825) (Moravia), and Camptopus lateralis (Germar, 1817) (Moravia); Bohemia: Sigara iactans Jansson, 1983; Moravia: Pachycoleus waltli Fieber, 1860, Aradus kuthyi Horváth, 1899; and Slovakia: Microvelia pygmaea, Heterocordylus cytisi Josifov, 1958, Aradus bimaculatus Reuter, 1872. Additional records of 22 rare species or species previously considered as regionally extinct in the Czech Republic and Slovakia are provided: Arctocorisa germari germari (Fieber, 1848), Notonecta reuteri reuteri Hungerford, 1928, Microvelia buenoi Drake, 1920, Salda henschi (Reuter, 1891), Salda muelleri (Gmelin, 1790), Agramma atricapillum (Spinola, 1837), Phytocoris hirsutulus Flor, 1861, Polymerus brevicornis (Reuter, 1879), Hyoidea notaticeps Reuter, 1876, Conostethus hungaricus Wagner, 1941, Harpocera hellenica Reuter, 1876, Peirates hybridus (Scopoli, 1763), Aradus crenaticollis R. Sahlberg, 1848, Aradus mirus Bergroth, 1894, Aradus signaticornis R. Sahlberg, 1848, Arocatus melanocephalus (Fabricius, 1798), Melanocoryphus albomaculatus (Goeze, 1778), Melanocoryphus tristrami (Douglas et Scott, 1868), Tropidophlebia costalis (Herrich-Schaeffer, 1850), Coriomeris scabricornis scabricornis (Panzer, 1805), Aelia rostrata Boheman, 1852, and Eurydema fieberi Fieber, 1837. The species Conostethus griseus Douglas et Scott, 1870 is excluded from the list of Slovak fauna. Including the new records presented in this paper, the number of true bug species recorded reaches 869 in the Czech Republic (762 in Bohemia and 810 in Moravia) and 850 in Slovakia. Thanks to the discovery of Tupiocoris rhododendri and Tuponia macedonica, the number of alien species of Heteroptera established in the Czech Republic increases to 22. Keywords. Heteroptera, true bugs, faunistics, distribution, biology, central Europe, Czech Republic, Slovakia 495 P. K MENT ET AL.

Introduction The present paper is a further continuation of the “New and interesting records of true bugs (Heteroptera) from the Czech Republic and Slovakia” series (KMENT & BRYJA 2001; KMENT et al. 2003, 2005; HRADIL et al. 2008). Since the last part of the series, faunistic research into Czech and Slovak Heteroptera has been far from idle, and a number of contributions presenting new records or regional faunal inventories have been published. The following new species have been added to the fauna of the Czech Republic: Corixa panzeri Fieber, 1848 (Corixidae) (recorded in Moravia, STRAKA et al. 2009), Lasiacantha hermani Vásárhelyi, 1977 (Tingidae) (Moravia, MALENOVSKÝ et al. 2011), Dicyphus escalerae Lindberg, 1934 (Miridae) (Moravia, HRADIL 2010), Charagochilus spiralifer Kerzhner, 1988, Heterocordylus cytisi Josifov, 1958 (both Miridae) (Moravia, KMENT & BAÒAØ 2012), Dysepicritus rufescens (A. Costa, 1847) (Anthocoridae) (Moravia, KMENT & BAÒAØ 2012), Nagusta goedelii (Kolenati, 1857) (Reduviidae) (Bohemia, KMENT & DOLEJŠOVÁ 2010), and Belonochilus numenius (Say, 1831) (Moravia, HRADIL 2011). Moreover, SYCHRA & KMENT (2009) have confirmed the occurrence of Sigara scotti (Douglas et Scott, 1868) (Corixidae) based on material from westernmost Bohemia, previously listed without exact records by ŠTUSÁK (1980). The fauna of Bohemia has been enriched with records of Cimex pipistrelli Jenyns, 1839 (Cimicidae) (BALVÍN et al. 2012), Tropidothorax leucopterus (Goeze, 1778) (Lygaeidae) (KMENT et al. 2009a) and Oxycarenus lavaterae (Fabricius, 1787) () (KMENT et al. 2010), and the Moravian fauna by records of Gerris lateralis Schummel, 1832 (Gerridae) (JEZIORSKI et al. 2013), Salda muelleri (Gmelin, 1790) (Saldidae) (ROHÁÈOVÁ 2011), Acalypta platycheila (Fieber, 1844) (Tingidae), Phytocoris hirsutulus Flor, 1861 (Miridae) (both KMENT & BAÒAØ 2012), Halticus saltator (Geoffroy, 1785) (Miridae) (HRADIL et al. 2013), Loricula ruficeps (Reuter, 1884) (), and Elatophilus nigricornis (Zetterstedt, 1838) (Anthocoridae) (both KMENT & BAÒAØ 2012). Among these additions, Dicyphus escalerae and Belonochilus numenius represent new alien species (HRADIL 2010, 2011), while the single recorded specimen of Nagusta goedelii found in synanthropic conditions in Prague is probably only a case of accidental introduction (KMENT & DOLEJŠOVÁ 2010). On the other hand, Heterocordylus leptocerus (Kirschbaum, 1856) (Miridae) has been excluded from the Moravian fauna due to a misidentification of H. cytisi (KMENT & BAÒAØ 2012). As well as the new records, two species previously considered extinct in the Czech Republic (KMENT & VILÍMOVÁ 2006) have recently been recorded again – Eurydema fieberi Fieber, 1837 (Pentatomidae) (Bohemia, KMENT & JINDRA 2008) and Pithanus hrabei Stehlík, 1952 in its terra typica in the Jeseníky Mts. (Moravia; LACINA 2010, 2011). A number of recent faunal inventories are relevant and worthy of mention. In Bohemia, the detailed results of long-term studies of aquatic insects in the mountain lakes of the Šumava National Park, also including aquatic and semi-aquatic true bugs, has been published by SOLDÁN et al. (2012). Further inventories covered the fauna of Pøebuz environs in the Krušné hory Mts (SYCHRA & KMENT 2009, MALENOVSKÝ et al. in press), the Èeský les Mts. (KMENT & KEJVAL 2011), the Pamferova huť locality in the Šumava National Park (DVOØÁK et al. 2010) and the Žïár Nature Park in south-western Bohemia

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(ZÁHLAVOVÁ 2009). Data on water bugs also appear in the summary of a long-term study of benthic macroinvertebrates of the River Labe (= Elbe) (KOLAØÍKOVÁ et al. in press). In Moravia, KMENT & BAÒAØ (2012) presented the results of extensive sampling in the Bílé Karpaty Protected Landscape Area, where 500 species were documented, thus promoting the Bílé Karpaty to the ranks of the best-explored areas in the Czech Republic. MALENOVSKÝ et al. (2011) presented an inventory of 15 well-preserved grassland sites in the Pannonian part of southern Moravia. Further, ROHÁÈOVÁ (2011, 2012a, b, 2013) published inventories of four reserves in the area of the Moravskoslezské Beskydy Mts. and the Javorníky Mts., while B. LIS (2012) made an inventory of the true bugs in the Štramberk Botanical Garden. In addition to locality inventories, detailed data for several species have been reviewed: Macrosaldula scotica (Curtis, 1835) (Saldidae) (KRIST & KMENT 2010), Cimex lectularius Linnaeus, 1758 (SADÍLEK et al. 2013), Cimex pipistrelli Jenyns, 1839 (BALVÍN et al. 2012), Tropidothorax leucopterus (KMENT et al. 2009a), Eurydema fieberi (KMENT & JINDRA 2008), and Pinthaeus sanguinipes (Fabricius, 1781) (Pentatomidae) (KMENT et al. 2009b), the last five of which also include records from Slovakia. Additional faunistic records are included in certain ecological papers published recently (e.g. ROHÁÈOVÁ 2008a, b; ROHÁÈOVÁ & DROZD 2009; SPITZER et al. 2010; SUCHÁ 2010; TROPEK et al. 2010, 2012; HRADIL et al. 2013). Considerably less attention has been paid to the fauna of Heteroptera in Slovakia than that of the Czech Republic. KMENT & BAÒAØ (2010) excluded Maccevethus corsicus corsicus Signoret, 1862 (Rhopalidae) from the Slovak fauna due to a misidentification of Maccevethus caucasicus (Kolenati, 1845), while KMENT & BAÒAØ (2012) corrected Slovak records of Charagochilus weberi Wagner, 1953 (Miridae) as belonging to misidentified Ch. spiralifer. KMENT & CUNEV (in press) submitted a manuscript to Entomofauna Carpathica in 2011 providing the first record of Belonochilus numenius from Slovakia; however, this has not yet been published. Not a single faunal inventory was published in Slovakia, with records of Heteroptera limited to incidental notes included in otherwise-focused papers (see BARTA 2008, 2009; KMENT & JINDRA 2008; KMENT et al. 2009a, b; KMENT & BAÒAØ 2010; KMENT & VILÍMOVÁ 2010; KOLLÁR et al. 2009; VINOKUROV 2009; MANKO 2011; BALVÍN et al. 2012; KRIŠTOFÍK & DANKO 2012; MATOCQ & PLUOT-SIGWALT 2012; VOTÝPKA et al. 2012; KNYSHOV & KONSTANTINOV 2013; SADÍLEK et al. 2013; ZHAO et al. 2013). The most promising event in Slovak Heteroptera research is the current species inventory and ecological study of aquatic and semi-aquatic Heteroptera (KLEMENTOVÁ 2012), of which only preliminary records have been published to date, including first Slovak records of Anisops sardeus Herrich- Schaeffer, 1849, Notonecta lutea Müller, 1776, Notonecta maculata Fabricius, 1794 (), Arctocorisa carinata C. R. Sahlberg, 1819, and Sigara distincta Fieber, 1848 (KLEMENTOVÁ et al. 2011, 2012a, b; KLEMENTOVÁ & SVITOK in press). A number of additional changes to the faunal lists of the Czech Republic have resulted from recent progress in heteropteran taxonomy. These include reinstating Maccevethus errans caucasicus (Kolenati, 1845) (Rhopalidae) to species status (KMENT & BAÒAØ 2010), synonymization of Amblytylus macedonicus Wagner, 1956 with Megalocoleus naso (Reuter, 1879) (MATOCQ & PLUOT-SIGWALT 2012), reinstatement of

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Plagiognathus (Plagiognathus) vitellinus (Scholz, 1847) to the resurrected genus Parapsallus Wagner, 1952, as Parapsallus vitellinus (Scholtz, 1847) (DUWAL et al. 2013), and synonymization of Ventocoris (Ventocoris) trigonus (Krynicki, 1871) with V. (V.) rusticus (Fabricius, 1781) (DURSUN & FENT 2013). The status of bat bugs of the Cimex pipistrelli species-group (i.e. C. pipistrelli, C. dissimilis (Horváth, 1910), and C. stadleri Horváth, 1935) in the western Palaearctic remains controversial, since neither morphometric nor DNA characters allow more than one taxon to be distinguished. BALVÍN et al. (2013) therefore suggested “treating C. dissimilis and C. stadleri as junior synonyms of C. pipistrelli until new evidence is available”. One additional change concerning our fauna, reinstating the subspecies Graphosoma (Graphosoma) lineatum italicum (O. F. Müller, 1766) (PÉRICART 2010), was not generally accepted (see e.g. AUKEMA et al. 2013). Including the new records presented in this paper, the number of the recorded true bug species reaches 869 in the Czech Republic (762 in Bohemia and 810 in Moravia) and 850 in Slovakia. Findings of Tupiocoris rhododendri and Tuponia macedonica bring the number of alien species of Heteroptera established in the Czech Republic to 22 (see also KMENT 2006; HRADIL et al. 2008; KMENT & DOLEJŠOVÁ 2010; RABITSCH 2010a; HRADIL 2010, 2011).

Material and methods The Czech Republic is here divided into Bohemia and Moravia, defined by their historical boundaries (see KMENT 2009). Codes from the Central European mapping grid (EHRENDORFER & HAMANN 1965) follow PRUNER & MÍKA (1996) and NOVÁK (1989). Our taxonomic system is after CASSIS & GROSS (1995, 2002) and the nomenclature follows AUKEMA & RIEGER (1995, 1996, 1999, 2001, 2006) and AUKEMA et al. (2013). The following abbreviations are used throughout the text: bor. – northern, centr. – central, mer. – southern, occ. – western, or. – eastern; ditto – at the same locality; NM – Nature Monument, NNM – National Nature Monument, NNR – National Nature Reserve, NP – National Park, NR – Nature Reserve, PLA – Protected Landscape Area, SAC – Special Area of Conservation (defined in the European Union’s Habitats Directive 92/43/EEC); spec. – unsexed imago; L – larva; ap – apterous, ma – macropterous; YPT – yellow pan trap. The material examined is deposited in the following collections: BMFC ...... Muzeum Beskyd [= Beskydy Museum], Frýdek-Místek, Czech Republic JCNS ...... Jozef Cunev collection, Nitra, Slovakia JSBC ...... Jan Sychra collection, Brno, Czech Republic JVPC ...... Jitka Vilímová collection, Praha, Czech Republic KHMC ...... Karel Hradil collection, Miletín, Czech Republic MCHD . . Muzeum Chodska [= Museum of Chodsko Region], Domažlice, Czech Republic MHBC ...... Michal Horsák collection, Brno, Czech Republic MMBC ...... Moravian Museum, Brno, Czech Republic MMHC ...... Marion Mantiè collection, Hluèín, Czech Republic MMLC ...... Municipal Museum Mariánské Láznì, Czech Republic MSHC ...... Michal Straka collection, Hustopeèe, Czech Republic

498 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

MVBC ...... Martin Vašíèek, Brno, Czech Republic NMPC ...... National Museum, Praha, Czech Republic OBPC ...... Ondøej Balvín collection, Praha, Czech Republic OKZC ...... Ondøej Konvièka collection, Zlín, Czech Republic OMOC ...... Ostravské muzeum [Museum of Ostrava], Ostrava, Czech Republic PKBC ...... Petr Komzák collection, Brno, Czech Republic TDCC ...... Tomáš Ditrich collection, Èeské Budìjovice, Czech Republic VHNC ...... Václav Hanzlík, Neratovice, Czech Republic VMPC ...... Východoèeské muzeum v Pardubicích [Museum of Eastern Bohemia in Pardubice], Pardubice, Czech Republic VZPC ...... Vladimír Zeman collection, Pøerov, Czech Republic ZJPC ...... Zdenìk Jindra collection, Praha, Czech Republic

Results Pachycoleus waltli Fieber, 1860 Material examined. Moravia centr.: Moravský kras PLA, Adamov env., under the Býèí skála NNR (49°18′25.258″N 16°41′37.042″E, 6666), alluvium along stream, sifting of wet moss covering dead branches with subsequent extraction using Winkler apparatus, 29.iii.2010, 3 ♂♂ 4 ♀♀ (one pair collected in copula), P. Baòaø lgt. et det., D. Rédei revid. (MMBC). Remarks. European species distributed from Ireland and Great Britain in the west to European Russia in the east, and from Sweden and Finland in the north to southern France and Bulgaria in the south (KERZHNER 1995, HEISS & PÉRICART 2007, AUKEMA et al. 2013). It lives in shaded habitats in permanently wet to very wet moss (Sphagnum, Hypnum, Brachythecium, Cratoneurum), i.e in various types of swamp, spring fen and wet meadow, seepages in forests, as well as on the shores of bodies of water and along small brooks and drains in more open situations; the specimens stay often fully submerged (AUKEMA et al. 2002, WACHMANN et al. 2006, HEISS & PÉRICART 2007). In the Czech Republic, the species was previously known only from the Sphagneto-Callunetum at Trstìnice (6042) in north-western Bohemia (ROUBAL 1957a, as P. rufescens). New species for Moravia.

CORIXIDAE Arctocorisa germari germari (Fieber, 1848) (Fig. 10)

Material examined. Bohemia bor.: Lomnice nad Popelkou (5458), v.1953, 1 ♂ 1 ♀, J. Roubal lgt. et det., P. Kment revid. (NMPC). Bohemia occ.: Pøebuz (5641), 4.viii.1980, 1 ♂, J. Pávek lgt., P. Kment det. (MMLC); according to the collector of the specimen, the precise locality may well be the Rolavský rybník (or Lieche) pond north of Pøebuz (910 m a.s.l.) (J. Pávek, pers. comm. to L. Dvoøák). Remarks. Euro-Siberian species, with the nominotypical subspecies distributed from Ireland and Norway to East Siberia; the other subspecies, A. g. mongolica Kanyukova, 2003, is endemic to Mongolia (JANSSON 1995, KANYUKOVA 2003, AUKEMA et al. 2013). Arctocorisa germari is a pelagic species inhabiting large oligotrophic waters, including those affected by acidification or slightly brackish ones, preferring deep waters with little

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 499 P. K MENT ET AL. or no vegetation and sandy bottom, especially lakes, dune lakes, fens, sand holes and sand pits (e.g. HENRIKSON & OSCARSON 1981, 1985; SAVAGE 1989; AUKEMA et al. 2002; WACHMANN et al. 2006). In Lusatia, the species was found in the lakes that form in open- pit coal mines, in the littoral and pelagic zones where the pH of the water was 2.2–3.8 and there were high concentrations of Fe and Al cations (WOLLMANN 1998, 2000). In the Czech Republic, the species is known from only four localities in Bohemia: TEYROVSKÝ (1930) collected two females in April 1928 in a reservoir several metres deep in a quarry near Babice (5954). In mid-May, 1953 ROUBAL (1957b) discovered a large population (40 ♂♂ 20 ♀♀) overwintering in a swimming pool in Lomnice nad Popelkou (5458), and he also reported a finding of 3 specimens in the Souš reservoir (51–5257) in the Jizerské hory Mts. Finally ŠTYS (1976) recorded a single female from the Fláje reservoir in the Krušné hory Mts. (5347), extracted from the stomach contents of an American brook trout (Salvelinus fontinalis (Mitchill, 1814)), together with two females of Arctocorisa carinata carinata (C. Sahlberg, 1819). Here we provide one additional record and confirm Roubal’s identification based on examination of male genitalia and strigil. However, recent extensive sampling of aquatic bugs in the environs of Pøebuz has not confirmed the occurrence of A. germari (see. SYCHRA & KMENT 2009, MALENOVSKÝ et al. in press).

Sigara iactans Jansson, 1983 Material examined. Bohemia bor.: Dìèín – Podskalí, Severní tùò pool (50°48′59″N 14°13′26″E, 5151), 14.ix.2011, 2 ♂♂, M. Straka lgt. et det., P. Kment revid. (MSHC, NMPC). Moravia bor.: Pustìjov, Pustìjovský potok brook (49°42′02.22″N 18°03′47.50″E; 6374), 24.vii.2002, 1 ♂, 22.ix.2002, 1 ♂, K. Petøivalská, P. Kment det. (NMPC); Pustìjovský potok brook (49°41′17.45″N 18°01′25.26″E; 6374), 28.x.2002, 2 ♂♂, K. Petøivalská, P. Kment det. (NMPC). Remarks. Ponto-Mediterranean species (distributed in Bulgaria, Greece, Macedonia, Romania, Russia (South European Territory), Turkey (both European and Asian), and Ukraine), established and spreading in north-western Europe (Belgium, Denmark, Czech Republic, England, France, Germany, the Netherlands, Poland, Sweden) (JANSSON 1995, AUKEMA et al. 2013). WACHMANN et al (2006) noted S. iactans from nutrient-poor, sparsely vegetated waters, partially with low pH; also as a pioneer species in newly- created reservoirs. In the Netherlands, most waters where S. iactans is found have little submerged vegetation and the acidity (pH) is generally alkaline (>7), the species avoids acid waters; inhabited waters are always permanent and usually quite large (eutrophic watercourses, mesothrophic sandpits); slow current is tolerated, but the species was only infrequently found in brooks. Strikingly often it is found in fresh and slightly brackish dune lakes on the West Frisian Islands and the species also occurs in “boezem” waters (= system of reservoirs of superfluous polder-water), but peat areas are not inhabited (CUPPEN 1988, AUKEMA et al. 2002). It has been collected in sandpits in England (NAU & BROOKE 2006), and in northern France in a pond with littoral vegetation (ELDER & CHÉRAU 2003). In the Czech Republic, it was previously recorded only from Poodøí PLA (6274–75) in northern Moravia, collected in eutrophic ponds with littoral vegetation and

500 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V an oxbow lake (BRYJA & KMENT 2001). KMENT & SMÉKAL (2002) later provided additional records from Poodøí PLA (the specimens from River Odra appear to have originated from a pond discharged few days ago) and one male from Slezské Pavlovice (Velký Pavlovický rybník Pond, 5672), also in northern Moravia. New species for Bohemia.

NOTONECTIDAE Notonecta (Notonecta) reuteri reuteri Hungerford, 1928 Material examined. Moravia bor.: Rejvíz, Rejvíz NNR (50°13′11.6″N 17°17′12.7″E, 5769), peat-bog, Velké mechové jezírko Pool, 768 m a.s.l., 15.viii.2011, 2 ♂♂ 1 ♀, P. Kment & J. Sychra lgt. et det. (JSBC, NMPC). Remarks. Euro-Siberian boreo-montane species with the nominotypical subspecies distributed from Ireland and Norway to Japan; N. r. ribauti Poisson, 1935 is confined to the Western Alps (POLHEMUS 1995). In the Czech Republic, the species is classified as endangered (KMENT & VILÍMOVÁ 2006). It usually lives in more or less oligothrophic fens on sandy soils, preferring complex habitats with abundant plants and dead plant matter; tyrphophilous with strong preference to peat bogs (SVENSSON et al. 2000, AUKEMA et al. 2002, WACHMANN et al. 2006). In the Czech Republic known largely from peat-bogs and vegetated dystrophic ponds in Bohemia (Soos NNR, Šumava NP, Novohradské hory Mts., Tøeboòsko PLA, and Èeskomoravská vysoèina Highlands (e.g. ŠTYS 1960, 1961; PAPÁÈEK 1991, 2002, 2004; KMENT & SMÉKAL 2002; SOLDÁN et al. 2012). However, in Moravia the species is known from only a single locality, the Velké mechové jezírko Pool in the Rejvíz NNR (TEYROVSKÝ 1950, STEHLÍK 1952, ŠTYS 1960). ŠTYS (1960) collected it there in 1955, together with the morphologically very similar N. lutea. KMENT & SMÉKAL (2002) recorded only N. lutea in the Velké mechové jezírko Pool. Here we confirm the occurrence of N. reuteri in Rejvíz after an interval of 57 years.

VELIIDAE Microvelia (Microvelia) buenoi Drake, 1920 (Figs 1, 11–14) Material examined. Bohemia centr.: Kokoøín, Kokoøínský dùl NR, Kaèírek pond (50°25′54.8″N 14°34′55.8″E, 5553), 24.viii.2009, 2 ♂♂ (ma) 2 ♂♂ (ap), P. Kment & J. Sychra lgt. et det. (JSBC); Chudíø, travertine spring fen above Olšový rybník pond (50°18′06.1″N 15°01′04.1″E, 5656), NE of Nový rybník pond, 235 m a.s.l., 21.viii.2012, 1 ♂ (ap), P. Kment & J. Sychra lgt. et det. (JSBC); Lysá nad Labem, Hrabanovská èernava NNR (50°12′49.7″N 14°49′38.1″E, 5754), shaded pool in SW corner of the reserve, 23.viii.2012, 1 ♂ (ma) 11 ♂♂ (ap) 1 ♀ (ma) 9 ♀♀ (ap), P. Kment & J. Sychra lgt., P. Kment det. (NMPC); Pøelouè, Labišťata, vegetated pools (50°2′37″N 15°32′24″E, 5959), 27.iv.2012, 1 ♂ (ap), M. Straka lgt. et det. (MSHC); Pøelouè, Støídeò, pool at road (50°2′47″N 15°34′34″E, 5959), 13.iv.2012, 1 ♂ (ap), M. Straka lgt. et det. (MSHC). Bohemia mer.: Horusice, Ruda NNR (49°09′03.9″N 14°41′27.7″E, 6854), 13.ix.2010, 1 ♀ (ap), P. Kment lgt. et det. (NMPC); Lužnice, Velký a Malý Tisý NNR, between Velký and Malý Dubovec ponds (49°03′58.4″N 14°43′22.0″E, 6954), shaded pool in Alnetum, 425 m a.s.l., 14.ix.2010, 1 ♂ (ma) 13 ♂♂ 3 ♀♀ (ap), P. Kment & J. Sychra lgt. et det. (JSBC, NMPC); Bukovec, Jizba pond (48°52′45.99″N 14°26′21.37″E, 7152), a single cove shaded by Alnetum, v.–ix.2011, 13 ♂♂ 16 ♀♀ (ap), v.–x.2012, 8 ♂♂ 10 ♀♀ (ap), 18.vi.2013, 2 ♂♂ 3 ♀♀ (ap), T. Ditrich lgt. et det. (TDCC). Moravia occ.: Ptáèov, Starý Ptáèovský rybník pond, littoral growth in NE part of the pond (49°13′39.25″N 15°55′20.01″E, 6761), 1.v.2011, 1 ♂ (ap), J. Sychra lgt. et det. (JSBC). Moravia mer.: Bojanovice, U Huberta SAC, abandoned fish-pond (48°57′11″N 16°00′00″E, 7061–62),

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19.viii.2009, 1 ♀ (ap), M. Straka lgt. et det., P. Kment revid. (MSHC); Ivaò, Betlém NR, pools (48°54′25″N 16°35′26″E, 7065) 25.v.2010, 1 ♂ (ap), M. Straka lgt. et det. (MSHC); Lednice, Pastvisko NNR, northern part (N 49°44′49.11″N 12°24′45.28″E, 7166), 26.iv.2003, 1 ♂ (ma), M. Horsák lgt., P. Kment det. (NMPC); Hrabìtice, Travní Dvùr SAC, reed pool (48°47′14″N 16°25′43″E, 7264), 30.iv.2009, 1 ♂ (ma) 2 ♂♂ (ap) 1 ♀ (ap), M. Straka lgt. et det. (MSHC); Hrabìtice, Travní dvùr SAC (48°47′19.8″N 16°25′41.0″E, 7264), wetland meadows, shaded ox-bow lake in a forest, 14.v.2011, 1 ♂ (ap), M. Horsák & P. Kment lgt., P. Kment det. (NMPC). Remarks. Holarctic species distributed from Great Britain to Central Asia and the Russian Far East, also in Canada and the USA (SMITH 1988, ANDERSEN 1995, AUKEMA et al. 2013). Minute species with two generations per year, overwintering as an adult; the majority of the specimens are apterous, with macropterous specimens occurring only rarely (WRÓBLEWSKI 1980). It is bound to quiet, shaded, exclusively standing waters, living in the littoral zones of lakes detached by stands of rushes and shaded by trees, in natural peaty waters (pools in peat-bogs), pools left by peat extraction, and small reservoirs in forests (especially within stands of alder). It is predatory, feeding mainly on collembolans (WRÓBLEWSKI 1980, WACHMANN et al. 2006). One exceptional find occurred on a shaded bank of the Podstola river in Poland, among grasses and leaf-litter (KRAJEWSKI 1969). KURZ¥TKOWSKA (1999) classified it as a grade II tyrphophilous species, inhabiting various dystrophic, polyhumic waters, often in forests. It is characterized in similar fashion by SAVAGE (1989) for the British Isles: rare, distributed in lakes, ponds and pools with a high content of organic matter in the bottom substrate, pH > 6 and conductivity 100–1000 µS/cm, at altitudes below 300 m. In Poland the species is considered widespread but rare (WRÓBLEWSKI 1980). In the Netherlands, AUKEMA et al. (2002) described the ecology of the species as follows: Microvelia buenoi occurs along the margins and shores of canals, ditches, fens and pools, where it can be especially abundant between emergent macrophytes and overhanging branches of shrubs and trees. On places without such vegetation, the species does not occur or is found only sporadically and in low numbers. It prefers a better water quality than M. reticulata (Burmesiter, 1835). In peat-moor areas it prefers a higher acidity than the latter and in the river area it prefers relatively natural habitats as ditches with Carex vegetation and original swamps. In cultivated landscape it is very rare (AUKEMA et al. 2002). Our recently-discovered localities are appropriate to these descriptions, including pools in alder stands and floodplain forests, small pools in peat-bogs and the shaded coves of ponds; the most interesting finding comes from a travertine spring fen, suggesting the species is not bound exclusively to low pH. In some localities M. buenoi occurred together with the common M. reticulata (e.g. Horusice, Ruda NNR; Chudíø, travertine spring fen; Lysá nad Labem, Hrabanovská èernava NNR; Bukovec, Jizba Pond, where M. buenoi regularly constituted 5–30 % of Microvelia population in one shaded cove). At the Travní Dvùr SAC locality we discovered M. buenoi, M. pygmaea and M. reticulata within one area of floodplain forests and meadows, although we cannot confirm their occurrence in the same microhabitat. Microvelia buenoi is classified as “endangered” in the Red List of Czech Heteroptera (KMENT & VILÍMOVÁ 2006). In the Czech Republic, the species was listed from Bohemia by HOBERLANDT (1977a, as M. umbricola Wróblewski, without any exact locality. ŠTUSÁK (1980) considered the

502 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V occurrence of the species in Czechoslovakia as unconfirmed. KMENT (1999) published first records for it from the Pálava PLA in southern Moravia (Pøítluky, Had, 7166; Mikulov, Nový rybník NR, 7266) (see also KMENT & SMÉKAL 2002). The first exact records for Bohemia were provided by KMENT & SMÉKAL (2002) and BRYJA & KMENT (2006a) in Kokoøínsko PLA (four localities in Kokoøínský dùl NR, 5553). Here we provide additional localities from both Bohemia and Moravia.

Microvelia (Picaultia) pygmaea (Dufour, 1833) (Figs 1, 15–17) Material examined. Moravia mer.: Ivaò, Nové Mlýny – Støední nádrž dam (48°54′34″N 16°36′12″E, 7065), 27.iv.2010, 1 ♀ (ap), M. Straka lgt. et det. (NMPC); Hrabìtice, Travní Dvùr SAC, slowly running canal (48°46′43″N 16°26′9″E, 7264), 12.vi.2009, 1 ♀ (ma), M. Straka lgt. et det. (MSHC); Úvaly, Úvalský rybník pond (48°44′45,6″N 16°42′42,4″E, 7266), 12.ix.2009, 1 ♂ (ap), J. Sychra lgt. et det., P. Kment revid. (JSBC). Slovakia occ.: Most pri Bratislave, Zelená voda gravel pit (48°9.234′N 17°15.083′E, 7869), recently abandoned gravel pit with sunlit shores, mineral bottom substrate and only sparse vegetation and detritus, 5.v.2011, 1 ♂ 3 ♀♀ (ap), M. Straka lgt. et det. (MSHC). Remarks. Holomediterranean–Central Asiatic species distributed from Great Britain and Morocco in the west to Kyrgyzstan and Tajikistan in the east, reaching as far north as the Netherlands, Germany, Austria, and Hungary (ANDERSEN 1995, HOFFMANN & MELBER 2003, AUKEMA et al. 2013). Microvelia pygmaea is a rare although quite widely- distributed species in the Pannonian lowland in Hungary (BENEDEK 1970; KONDOROSY et al. 1996; KONDOROSY & FÖLDESSY 1998; JUHÁSZ et al. 1999; KONDOROSY 2000, 2001, 2011; BAKONYI et al. 2002; BODA et al. 2004, 2012; VÁSÁRHELYI & BÁKONYI 2005; VÁSÁRHELYI et al. 2005, 2007; BODA 2006; NOSEK et al. 2007; SOÓS et al. 2009; KÁLMÁN et al. 2011; MÓRA et al. 2011) and Romania (DAVIDEANU 1999). The only verified record of this species in Austria is from historical material collected in Vorarlberg (close to the Swiss border) in the 1920s (RABITSCH 1999a). WACHMANN et al. (2006) characterized its habitat as sheltered, vegetation-rich standing waters with reed belts along the shore; they reported it from oxbow lakes surrounded with trees, as well as from swamps and ditches in meadows. In Hungary, BAKONYI et al. (2002) collected the species in a ditch by the road, and BODA et al. (2004) in stands of Carex in a marsh. In the British Isles, the species is limited to southern England and altitudes < 300 m; SAVAGE (1989) summarized its ecological requirements as standing waters (more frequently lakes than pools and ponds), its frequency growing with increasing plant cover and content of organic matter in the bottom substrate, conductivity < 1000 µS/cm. In the Netherlands, M. pygmaea occurs along the margins of eutrophic canals, canalized brooks and pools, being especially numerous close to the shores between emergent macrophytes and overhanging branches of shrubs and trees. After disturbance they move to open water. The species is only rarely found along shores without vegetation or with only a small strip of emergent vegetation, and if so, only in low numbers. It is only occasionally found in acid, running or temporary waters (AUKEMA et al. 2002). There are far fewer macropterous specimens than apterous within a given population (WACHMANN et al. 2006). For identification of Microvelia pygmaea and the two remaining European Microvelia species, see SAVAGE (1989) and KANYUKOVA (2006). New species for the Czech Republic and Slovakia.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 503 P. K MENT ET AL.

Fig. 1. Distribution of Microvelia buenoi Drake, 1920 and M. pygmaea (Dufour, 1833) in the Czech Republic. – occurrence of M. buenoi only, – occurrence of both M. buenoi and M. pygmaea.

SALDIDAE Salda henschi (Reuter, 1891) (Figs 2, 18) Material examined. Moravia bor.: Rejvíz, Rejvíz NNR (50°13′11.6″N 17°17′12.7″E, 5769), 21.iv.1993, 1 spec., I. Malenovský lgt., P. Kment det. (NMPC); ditto, Velké mechové jezírko, in wet Sphagnum at shores of the pool, 768 m a.s.l., 15.viii.2011, 2 ♂♂, P. Kment, V. Syrovátka & J. Sychra lgt. et det. (NMPC). Remarks. European boreo-montane species known from southern Sweden, Hogland Island in the Gulf of Finland (North European Territory of Russia), the Alps and other mountains of central Europe (Austria, Czech Republic, Germany, Liechtenstein, Poland, Slovakia, and Switzerland) (HOBERLANDT 1977b; LINDSKOG 1991, 1995). It usually lives in very wet Sphagnum growths in raised bogs (LINDSKOG 1991, WACHMANN et al. 2006). In the Vysoké Tatry Mts. [= High Tatras] (Slovakia), it was also collected on the clayey bank of a small mountain brook in a coniferous zone (HOBERLANDT 1977b). The species is classified as endangered in the Czech Republic (KMENT & VILÍMOVÁ 2006). In Bohemia, it was recorded from the Šumava Mts. (Poleènice, 7150), the Orlické hory Mts. (Jelení lázeò NR at Šerlich, 5664) and the Krušné hory Mts. (Boží Dar, 5543) (HOBERLANDT 1977b). In Moravia, the species is known only from the Velké mechové jezírko Pool near Rejvíz (Jeseníky Mts.), with published records for 1946–1964 (STEHLÍK

504 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

Fig. 2. Distribution of Salda muelleri (Gmelin, 1790) () and S. henschi (Reuter, 1891) () in the Czech Republic.

1952, HOBERLANDT 1977b). Here we confirm the occurrence of the species in the Jeseníky Mts.

Salda muelleri (Gmelin, 1790) (Fig. 2) Material examined. Bohemia bor.: Hrabìtice, Tichá øíèka NR (5257), 750 m a.s.l., 30.vii.–29.viii.2004, YPT, 1 ♂ 1 ♀, J. Preisler & P. Vonièka lgt., K. Hradil lgt. et det. (KHMC); Jizerské hory Mts. (no details), 8.–15.viii.1981, 1 ♂, P. Bílek lgt., Z. Jindra det. (ZJPC); Jizerské hory Mts., Horní Maxov, Malá Strana NR (5257), 20.vi.2009, 1 ♀, K. Hradil lgt. et det. (KHMC); Krkonoše Mts., Špindlerovka (5259), 26.vii.2000, 1 ♂, K. Hradil lgt. et det. (KHMC); Fláje env. (5347), 13.vi.1999, 4 ♂♂, J. Vilímová lgt., P. Kment det. (NMPC); Hora Svatého Šebestiána env. (5445), 18.x.1983, pitfall trap, 1 ♀, K. Absolon lgt., P. Kment det. (NMPC). Bohemia occ.: Pøebuz, Rotava river (5641), 12.vii.1990, 2 ♀♀, V. Šilha lgt., Z. Jindra det. (ZJPC); Kateøina, NPR Soos (5840), NPR Soos (5840), 8.vi.1975, 1 ♀, 23.vi.1978, 2 ♀♀, J. Strejèek lgt., Z. Jindra det. (ZJPC); ditto, 11.–12.vii.1989, 2 ♀♀, J. Vilímová lgt., P. Kment det. (NMPC). Bohemia mer.: Èerná v Pošumaví, Slavkovice env. (7250), 23.vi.1986, 1 ♂ 1 ♀, Z. Jindra lgt. et det. (ZJPC). Moravia bor. or: Bukovec, Bukovec NR (49°32′57.27.5′′N 18°51′31.6′′E, 6479), 500 m a.s.l., 29.v.-19.vi.2013, pitfall trap in the most swampy part of the mire meadow (Calthion without Sphagnum spp. tufts), 3 larvae; ditto, 19.vi.–30.vii.2013, 4 ♀♀, together with numerous specimens of Chartoscirta cocksii, M. Roháèová lgt. et det. (BMFC). Remarks. Euro-Siberian, boreo-montane species distributed from Ireland and France to the Far East of Russia (LINDSKOG 1995, AUKEMA et al. 2013). In the Czech Republic, the

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 505 P. K MENT ET AL. species is confined to peat-bogs, where it lives in wet tufts of Sphagnum spp. (HOBERLANDT 1977b), but in the Soos NNR it was also found in a salt marsh on bare soil under rushes (ROUBAL 1957b, ŠTYS 1961). In other parts of its area of distribution it also occurs in salt marshes along sea coasts (AUKEMA et al. 2002, WACHMANN et al. 2006). In the Czech Republic, it was previously known from only a few localities in Bohemia: Wies (= Stráž u Chebu, village abandoned after World War II; 5940) (DALLA TORRE 1877, as S. flavipes), Soos NNR (ROUBAL 1957a, ŠTYS 1961, HOBERLANDT 1977b), Krkonoše Mts. (former Lišèí bouda cottage in Lišèí louka Meadow, 5360) (ROUBAL 1959), the Šumava Mts. (Jezerní slať raised bog, 6947; Železná Ruda, 6845) (BOZDÌCHOVÁ 1973, HOBERLANDT 1977b), and the Krušné hory Mts. (Slatinný potok Brook, 5541; Chaloupky, 5641; Myší kout, 5641) (MALENOVSKÝ et al. in press). In Moravia, it was previously known from only a single locality – Obidová NM (6477) in the Moravskoslezské Beskydy Mts. (ROHÁÈOVÁ 2011). Here we provide additional localities for this endangered species (cf. KMENT & VILÍMOVÁ 2006).

TINGIDAE Agramma (Agramma) atricapillum (Spinola, 1837) (Fig. 19) Material examined. Morvia mer.: Hrabìtice, Travní dvùr SAC (48°47′19.8″N 16°25′41.0″E, 7264), wetland meadows, on Bolboschaenus maritimus, 14.v.2011, 10 ♂♂ 46 ♀♀, M. Horsák & P. Kment lgt., P. Kment det. (9 ♂♂ 42 ♀♀ in NMPC, 1 ♂ 4 ♀♀ in MMBC, 1 ♂ 7 ♀♀ in KHMC). Slovakia mer.: Hajnáèka, Velká dolina valley (7785–86), 20.v.2007, 1 ♀, J. Ch. Vávra lgt., P. Kment det. (OMOC); Pataš, Èiližské moèiare SAC (47°52′26.5″N 17°40′15.1″E, 8172), reed swamp near Èiližský potok brook, 12.v.2012, 1 ♀, P. Kment lgt. et det. (NMPC). Remarks. Holomediterranean species, extending to central Asia, known in central Europe from only the Pannonian lowlands (PÉRICART & GOLUB 1996, AUKEMA et al. 2013). Agramma atricapillum occurs in humid habitats, both saline and non-saline. It develops on various plants of the families Juncaceae, Cyperaceae and Typhaceae (Juncus, Scirpus, Bolboschoenus, Carex, Schoenoplectus, Typha) (PÉRICART 1983, WACHMANN et al. 2006). In Slovakia, it was first recorded from Moèenok (7773) and Gemerské Dechtáre (7786) by KMENT et al. (2003). In the Czech Republic, the species was previously known from only two specimens collected in a saline meadow near Vrbovec (7162) in southernmost Moravia (BALVÍN 2007). Both Moravian records are also in close proximity to the only two known localities in Lower Austria, which are Zwingendorf (RABITSCH 1999b) and Laa an der Thaya (W. Rabitsch, pers. comm.). Here we provide additional records for Moravia and Slovakia.

MIRIDAE Conostethus hungaricus Wagner, 1941

Material examined. Slovakia mer.: Kamenín (8177), 30.v.1953, 24 ♂♂ 34 ♀♀, Exc. M. N. Pragae lgt., P. Kment det. (NMPC); ditto, 6.vi.1960, 16 ♂♂ 18 ♀♀, Exc. M. N. Pragae lgt., L. Hoberlandt 1960 det. as C. salinus, P. Kment revid. (NMPC); Ve¾ké Kosihy, Mostová NR (= Derhidia) (8273), salt meadow, collected walking on the soil surface among sparse halophilous vegetation, 11.v.2012, 1 ♂ 3 ♀♀, P. Kment lgt. et det.

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(NMPC); Èenkov (8277), 7.vi.1960, 1 ♂, Exc. M. N. Pragae lgt., L. Hoberlandt 1960 det. as C. salinus, P. Kment revid. (NMPC). Remarks. Ponto-Mediterranean–Central Asiatic species distributed in the steppe zone from Austria to East Siberia, Mongolia and Uzbekistan (KERZHNER & JOSIFOV 1999, AUKEMA et al. 2013). Conostethus hungaricus is an inhabitant of salt marshes; WAGNER (1965) gave Lepidium crassifolium (Brassicaceae) as the host plant, but other authors suggest it probably lives on Poaceae as well (e.g. Agrostis stolonifera ssp. maritima, Puccinellia limosa) (see WAGNER 1941, MELBER et al. 1991, WACHMANN et al. 2004). In Slovakia, it has been previously recorded only from Èenkov (8277) (STEHLÍK & ŠTEPANOVIÈOVÁ 1961). Another species, Conostethus griseus Douglas et Scott, 1870, was recorded under its junior synonym C. salinus J. Sahlberg, 1870, from Kamenín (salt marsh, 6.vi.1960) (STEHLÍK 1961, HOBERLANDT 1963) and Èenkov (sands, 7.vi.1960) (HOBERLANDT 1963). According to the identification characters given by WAGNER (1952), particularly body length not exceeding 3.8 mm, all the Slovak specimens belong to the single species C. hungaricus. Further, it is possible that both C. griseus and C. hungaricus are just synonyms (see MATOCQ 1991). ŠTEPANOVIÈOVÁ & BIANCHI (2001) included C. hungaricus in the Red List of Slovak Heteroptera as an endangered species. Here we present a recent record of C. hungaricus from Slovakia, while C. griseus is excluded from the list of the Slovak fauna.

Harpocera hellenica Reuter, 1876 (Fig. 21) Material examined. Moravia mer.: Mohelno, Mohelenská hadcová step NNR (N 49°06′31.53″ E 16°11′12.99″, 6863), 11.v.2013, 2 ♀♀, M. Horsák, P. Kment & Biotým lgt. (NMPC). Remarks. Ponto-Mediterranean species recorded from Albania, Bulgaria, Greece, Asian Turkey, Israel and Jordan (KERZHNER & JOSIFOV 1999, CARAPEZZA 2002, AUKEMA et al. 2013), recently found extending into the Pannonian lowland (Czech Republic, Austria, possibly also Slovakia) (KMENT et al. 2005, RABITSCH 2010b). A zoophytophagous species living on oaks (Quercus spp.). It overwinters in the egg stage and adults appear for only a short period in May (JOSIFOV 1978, KMENT et al. 2005, RABITSCH 2010b). In the Czech Republic, it was first recorded on the basis of two females collected on Dìvín Hill (7065) on May 9, 2003. It was later found in Malaise trap samples from Machová NR near Javorník (7171c) (8.–28.v.2007) and Hryzlácké Mlýny near Nová Lhota (7171b) (7.iv.–12.v.2009) in the Bílé Karpaty Protected Landscape Area, and by pyrethroid spraying of apple trees in orchards at Velké Bílovice (7167) (11.v.2011) in southern Moravia. Here we present additional Moravian record from a well-sampled locality further west and north, suggesting a possible recent spread of the species within the Pannonian lowland.

Heterocordylus (Heterocordylus) cytisi Josifov, 1958 (Figs 3–7)

Material examined. Moravia occ.: Tøebíè (6761), no date, 1 ♂ 2 ♀♀, J. L. Stehlík lgt., P. Kment det. (MMBC); Studenec (6762), 5.vii.1942, 2 ♂♂, J. L. Stehlík lgt., P. Kment det. (MMBC); Bøezník, valley of

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Fig. 3. Distribution of Heterocordylus cytisi Josifov, 1958 in the Czech Republic.

Chvojnice river (6863), 28.vi.1942, 1 ♂, J. L. Stehlík lgt., P. Kment det. (MMBC); Mohelno, Mohelenská hadcová step NNR (6863), rocky steppe on serpentinite, western corner of the reserve, 350–400 m a.s.l., 20.vi.1975, 1 ♀, J. L. Stehlík lgt. (MMBC); Senorady (6863), v.1943, 1 ♀, Matoušek lgt., P. Kment det. (MMBC). Moravia centr.: Žerùvky, Baba hill (ca. 4 km N of Olšany) [= Na skále NNM] (6469), secondary steppe on loess loam, 240–260 m a.s.l., 6.vii.1963, 1 ♀, P. Lauterer lgt., P. Kment det. (MMBC); Kozárov (6564), meadows and forest undergrowth between Rašov and Kozárov, 600–630 m a.s.l., 3.vii.1970, 2 ♂♂ 1 ♀, L. Pospíšilová lgt. (MMBC); Zdounky, Divoky (6769), xerothermic vegetation (Inuletum ensifoliae), 10.vii.1963, 1 ♂, L. Pospíšilová lgt. (MMBC); Jedovnice, Rakovecké údolí valley (6666–67, 6766–67), meadow in forest, 11.vii.1969, 1 ♂, J. L. Stehlík lgt. (MMBC). Moravia mer.: Øíèany, slopes above mill (6764), steppe, 26.vi.1963, 1 ♂ 1 ♀, L. Pospíšilová lgt. (MMBC); Brno – Medlánky, Medlánecká skalka NM (49°14′29″N 16°34′03″E, 6765c), 275–300 m a.s.l., 26.v.2012, 1 ♂ 1 ♀, I. Malenovský lgt. (NMPC); Kývalka, 1.5 km W towards Omice (6864), forest margins on granodiorite, 380 m a.s.l., 1.vi.1964, 2 ♂♂ (newly hatched), P. Lauterer lgt. (MMBC); Kurdìjov, Nová hora hill (6966), steppe on loess, 300–350 m a.s.l., 8.vii.1971, 1 ♀, J. L. Stehlík lgt., P. Kment det. (MMBC); Vranov nad Dyjí, along road to Lanèov, ca. 1 km W of the village (7060), forest margins and mesic to xerothermic meadows on schist, 400–415 m a.s.l., 28.vi.1976, 1 ♂ 1 ♀, J. L. Stehlík lgt., P. Kment det. (MMBC); Klentnice, Kotelná [= Kotel hill]–Klausen [= Soutìska] (7165), S margin of forest E of Kotelná hill, 350–370 m a.s.l., 10.vi.1964, 2 ♂♂ 3 ♀♀, P. Lauterer lgt., P. Kment det. (MMBC); Klentnice, Tabulová hora hill (7165), 25.v.1959, 2 ♂♂, J. L. Stehlík lgt., P. Kment det. (MMBC); Perná (towards Horní Vìstonice), SW–W slope of Kotelná hill NE of Perná (7165), steppe on limestone, 330–400 m a.s.l., 20.v.1982, 1 ♀, J. L. Stehlík lgt. (MMBC); Pavlov (7165–66), ruderal vegetation among fields and vineyards, on limestone, 270 m a.s.l., 4.vii.1974, 2 ♂♂ 2 ♀♀, J. L. Stehlík lgt., P. Kment det. (MMBC); Hlohovec, Stará hora hill [Kamenice u Hlohovce NM] (7266), steppe on sandy limestone, 214 m a.s.l., 25.v.1971, 1 ♂ 1 ♀, J. L. Stehlík lgt., P. Kment det. (MMBC); Valtice (7266), 15.vi.1963, 1 ♂ 1 ♀, V. Krejèí lgt. (MMBC). Slovakia bor.: Hornádská kotlina valley, Spišské Podhradie, Dreveník hill (7090), steppe and

508 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

Fig. 4. Distribution of Heterocordylus cytisi Josifov, 1958 in Slovakia.

pasture vegetation on limestone, 600 m a.s.l., 11.vi.1971, 1 ♀, J. L. Stehlík lgt. (MMBC). Slovakia occ.: Trenèín (70–7174), no date, 1 ♂ 2 ♀♀, Èepelák lgt. (NMPC); Šaštín, È.[= Èervený] rybník NNR (7368), 29.v.1950, 2 ♀♀, J. L. Stehlík lgt. (MMBC); Malé Karpaty Mts., Smolenice (74–7570), 2.vii.1962, 1 ♂, V. Krejèí lgt. (MMBC); Bratislava (7868), no date, 2 ♂♂, no collector (NMPC). Slovakia centr.: Bojnice, Kalvaria hill (7277), forest steppe and forest margins on limestone, 350–450 m a.s.l., 21.vi.1964, 2 ♂♂, P. Lauterer lgt. (MMBC). Slovakia or.: Vyšné Raslavice (6893), pasture SW of the village, xerothermic vegetation, 320 m a.s.l., 19.vi.1965, 1 ♂ 1 ♀, L. Pospíšilová lgt. (MMBC). Slovakia mer.: Štiavnické pohorie Mts., Èajkov, above Èajkovský potok brook (76–7777), steppe and forest steppe on andesite, 190–300 m a.s.l., 5.vi.1971, 1 ♂, J. L. Stehlík lgt. (MMBC); Pohronská pahorkatina Hills, Mochovce, Dembrica [= Dobrica] hill (7776), rocky steppe and undergrowth of sparse forest on andesite slopes, 280–320 m a.s.l., 8.vi.1971, 1 ♀, L. Pospíšilová lgt. (MMBC). All P. Kment det.

Remarks. Balkan-Pannonian species described from Bulgaria by JOSIFOV (1958) and later recorded from Transcarpathian Ukraine (ROSHKO 1976), Macedonia (JOSIFOV 1986, without exact locality, see PROTIÆ 1998), Slovenia (GOGALA & GOGALA 1986, GOGALA 2006), and southern Moravia in the Czech Republic (KMENT & BAÒAØ 2012). In Moravia it had previously been recorded from Kobylí (7167) (HOBERLANDT 1947, as H. leptocerus) and Svatý kopeèek NR near Mikulov (7165) (BRYJA & KMENT 2006b, as H. genistae), Malhostovická pecka NM near Malhostovice (6664), and several localities in the southern part of the Bílé Karpaty PLA: Radìjov: Žerotín NM (7169b), Holý vrch Hill (7169–70); Knìždub, Èertoryje NNR (7170); Javorník: loam-pit (7171a), Machová NR (7171), SW slopes of Háj hill (7171a), Výzkum Hill (7171c); Velká nad Velièkou:

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 509 P. K MENT ET AL. towards Javorník (7171a); Nová Lhota: Fojtické Mlýny (7171b) (KMENT & BAÒAØ 2012). Recently, P. Kment examined several hundred Heterocordylus specimens, collected in the territory of the former Czechoslovakia, preserved in the collections of NMPC and MMBC. These yielded additional localities for H. cytisi as well as for the morphologically very similar H. genistae (Scopoli, 1763). Heterocordylus genistae, living on Genista spp. and Cytisus spp. (including C. scoparius) (WACHMANN et al. 2004) was found to be around ten times as numerous as H. cytisi (1131: 92 specimens) in the samples examined (in terms of both numbers of localities and specimens) and quite widely distributed in both thermophyticum and mesophyticum throughout Bohemia, Moravia and Slovakia. On the other hand, H. cytisi appears to be a rare species limited to central and southern Moravia and Slovakia, collected only in the thermophyticum. To judge by available locality data, H. cytisi is bound to xerothermic grassland localities, usually steppes, forest steppes and xerothermic meadows on hill-slopes, predominantly on basic substrates (limestone, andesite, loess, flysh sediments), although there are two records on acidic substrates as well (granodiorite, schist) (KMENT & BAÒAØ 2012, this paper). However, H. cytisi is locally more abundant than H. genistae (limestone hills of the Pavlovské kopce Hills, xerothermic species-rich meadows on basic flysh in the south of the Bílé Karpaty PLA). The localities for H. cytisi are situated mostly at altitudes of between 200 and 400 m, but in ideally suitable localities it has been recorded as high as 630 m (KMENT & BAÒAØ 2012, this paper). The occurrence of H. cytisi together with H. genistae on a same site has been documented only rarely, e.g. in the localities of the Chvojnice river valley (with 11 ♂♂ 13 ♀♀ of H. genistae), Kozárov (with 1 ♂ 1 ♀ of H. genistae) and Smolenice (with 1 ♀ of H. genistae) (this paper). In Moravia and Slovakia, adults of H. cytisi were collected between May 20 and July 11 (KMENT & BAÒAØ 2012, this paper). The only known host plant of H. cytisi is Chamaecytisus (= Cytisus) supinus (JOSIFOV 1958, 1963; GÖLLNER-SCHEIDING & ARNOLD 1988). According to the locality data from the MMBC database, “Cytisus sp.” was mentioned among plant species sampled in the localities of Bojnice, Kývalka, Vranov nad Dyjí, Vyšné Raslavice, and Žerùvky. The distribution of Ch. supinus in Moravia corresponds roughly with known records of H. cytisi (see SKALICKÁ 1995). However, a closer comparison shows that Ch. supinus does not grow in some of the localities, e.g. Kurdìjov, Kobylí, the Pavlovské kopce Hills, and the environs of Radìjov and Velká nad Velièkou in the Bílé Karpaty Mts., where it is replaced by the closely-related Ch. virescens (V. Grulich, pers. comm.). Heterocordylus cytisi is therefore definitely not a monophagous species, and its host- plant relationships will reward further investigation. Heterocordylus cytisi is absent from the basic identification keys for the Central European Heteroptera (e.g. WAGNER 1952, 1967; WAGNER & WEBER 1964) so it has remained overlooked, previously misidentified as either H. genistae (females) or H. leptocerus (Kirschbaum, 1856) (males). However, the males may easily be differentiated by the structure of the parameres, using the original description (JOSIFOV 1958) as well as the keys by KERZHNER (1964) and WAGNER (1974). For identification of females, we

510 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

Figs 5–9. Squama covering base of oviposior in females (ventral view). 5–7 – Heterocordylus cytisi Josifov, 1958, 8–9 – Heterocordylus genistae (Scopoli, 1763).

recommend the shape of the squama covering the base of the ovipositor, which is short, about equilaterally triangular in H. genistae (Figs 8–9), while it is elongated, in the form of a narrow isosceles triangle in H. cytisi (Figs 5–7) (see also KERZHNER 1964). Because of the discovery of H. cytisi, the occurrence of Heterocordylus leptocerus in Slovakia requires confirmation. In Slovakia, the species was recorded from Svätý Jur (BALTHASAR 1937) and recently from Devínska Kobyla Hill near Bratislava (HERCZEK & LUKÁŠ 1997). Unfortunatelly the voucher specimens of the latter record are lost (A. Herczek, pers. comm.) so we are currently unable to confirm occurrence of the species in Slovakia. Despite the voucher specimen of BALTHASAR’S (1937) record is supposed to be in Slovak National Museum in Bratislava we prefer to exclude H. leptocerus from the list of Slovak fauna for the time being.

Hyoidea notaticeps Reuter, 1876 (Fig. 20)

Material examined. Slovakia mer.: Èenkov (8277), 8.vi.1965, 58 ♂♂ 294 ♀♀ 1 L, M. Kocourek lgt., P. Kment det. (52 ♂♂ 213 ♀♀ 1 L in NMPC, 6 ♂♂ 81 ♀♀ in MMBC); Èenkov, Èenkovská step NNR (N 47°46′07.6″ E 18°31′12.0″, 8277), Stipa steppe on sands with intermixed Ephedra distachya, 13.v.2011, 1 ♂ 1 ♀, P. Kment lgt. et det. (NMPC). Remarks. Species of the Palaearctic steppes, distributed from Ukraine and the Southern Territory of European Russia further east to Iran, Tajikistan, and Inner Mongolia in northern China; there is also an isolated disjunct area of distribution in the Pannonian lowlands (southern Slovakia, Hungary) (KERZHNER & JOSIFOV 1999, AUKEMA et al. 2013, KNYSHOV & KONSTANTINOV 2013). The species is trophically bound to Ephedra species

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 511 P. K MENT ET AL.

(E. distachya, E. strobilacea, E. intermedia) (HOBERLANDT 1963, PUTSHKOV & PUTSHKOV 1983, KAPLIN 1993, KNYSHOV & KONSTANTINOV 2013). It is univoltine and larvae were collected from the end of March until the middle of May, adults from April until May in Turkmenistan (KAPLIN 1993), while in central Europe the adults occur from the middle of May until June (HOBERLANDT 1963). In Slovakia, the species was first recorded from “Èenkov” based on 31 ♂♂ and 40 ♀♀ collected between June 6 and 11 in 1960–1962 by L. Hoberlandt and P. Štys (HOBERLANDT 1963). These records refer, in fact, to the Èenkovský les Reserve, near Mužla, which is the single locality for its host plant – Ephedra disatchya – in Slovakia. ŠTYS (1972) reported another abundant population of E. distachya also hosting numerous H. notaticeps on the northern margin of Èenkovský les Forest c. 2–3 km from the reserve (5.–6.vi.1965, P. Štys & M. Kocourek lgt.). The numerous specimens labelled as “Èenkov” with the same date, deposited in the NMPC and MMBC collections originate from this sample. Besides the published records, P. Štys regularly observed (not collected) larvae and adults of this species on Ephedra distachya at this locality during intermittent visits between 1995 and 2010 (P. Štys, pers. comm.). KNYSHOV & KONSTANTINOV (2013) studied several specimens from Èenkov originating from samples published by HOBERLANDT (1963) and ŠTYS (1972), and 1 ♂ from Štúrovo (47.8°N 18.7333°E, 18.vi.1991, H. Günther lgt.). However, the record from Štúrovo lies, according to the geographical co-ordinates given, in Kamenica nad Hronom, where the host plant does not grow (V. Grulich, pers. comm.), strongly suggesting a mislabelling. ŠTYS & ŠKAPEC (1992) included H. notaticeps in the first edition of the Red List of Czechoslovakia as an “endangered species”, but provided no additional records. ŠTEPANOVIÈOVÁ & BIANCHI (2001) classified the species as “critically endangered” in Slovakia. Hyoidea notaticeps as well as its host plant, E. distachya, are remarkable examples of glacial relicts surviving in the remnants of the Pleistocene loess steppes as represented by the steppe localities of the Pannonian lowlands (BEZUS’KO 1999). Here we confirm recent occurrence for this species in Slovakia.

Isometopus mirificus Mulsant et Rey, 1879 (Fig. 22)

Material examined. Moravia mer.: Tvrdonice env., Rýnava (7268a), 27.vii.2012, 1 ♀, L. Bobot lgt., P. Kment det. (NMPC). The locality is an elevation in hardwood floodplain forest (Aesculus, Quercus, Tilia, Fraxinus); the species was collected at light (blacklight-blue bulb) during a warm and foggy night (over 20°C at midnight) (L. Bobot, pers. comm.). Remarks. Generally northern Mediterranean species, known from France, Italy, Germany, Hungary, Serbia, Romania, Bulgaria, Greece, Ukraine, and Asian Turkey (KERZHNER & JOSIFOV 1999, KONDOROSY 2005, PROTIÆ 2008, AUKEMA et al. 2013); listed in error for Iran by AUKEMA et al. (2013), based on a misinterpretation in LINNAVUORI et al. (1998). On the other hand, both KERZHNER & JOSIFOV (1999) and AUKEMA et al. (2013) overlooked recent records from Germany, where the species was discovered quite recently in Baden-Württemberg, Rhineland-Palatinate and Hesse (RIETSCHEL 2000, SIMON 2002, WACHMANN et al. 2004). It dwells on the bark of leafy trees (Juglans,

512 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

Quercus pubescens, Pyrus) (PÉRICART 1965, RIETSCHEL 2000, SIMON 2002, WACHMANN et al. 2004, PROTIÆ 2008). The larvae occur in May and July, adults appear from June to August and September, the females living longer (WACHMANN et al. 2004). New species for the Czech Republic.

Phytocoris (Phytocoris) hirsutulus Flor, 1861 Material examined. Moravia centr.: Brno – Nový Lískovec (6865), 305 m a.s.l., on Malus domestica, 21.vi.2012, 1 ♂, P. Šťastná lgt., K. Hradil det. (KHMC). The apple trees were fogged (Puls Fog) with an insecticide containing deltamethrin and the dead insects were collected from canvas placed under the crowns of the trees. Remarks. A very rare central European species distributed in Germany, Latvia (KERZHNER & JOSIFOV 1999), Slovenia (FLOREN & GOGALA 2002, GOGALA 2006), Estonia (COULIANOS 2005), Czech Republic (HRADIL et al. 2008), and Lithuania (Coulianos in AUKEMA et al. 2013). It dwells on the bark of old fruit trees, e.g. apple (Malus domestica) and pear (Pyrus communis), but is occasionally found on oak (Quercus) (SIMON 1992, MORKEL 2006, WACHMANN et al. 2004) and beech (Fagus sylvatica) as well (FLOREN & GOGALA 2002, GOGALA 2006). In the Czech Republic, it was first found in 2002 in Bohemia (Holovousy, 5659) (HRADIL et al. 2008), while in Moravia it was first encountered in Malaise trap samples from Jazevèí NNR (7171) in the Bílé Karpaty Mts. in 2009 (KMENT & BAÒAØ 2012). Second record for Moravia.

Polymerus (Poeciloscytus) brevicornis (Reuter, 1879)

Material examined. Slovakia mer.: Nitra, Kalvária hill and city (7774a), 140–273 m a.s.l., 19.vi.1984, 2 ♀♀, P. Lauterer lgt., J. Bryja det. (MMBC); Ve¾ké Kosihy, Mostová NR (= Derhidia) (8273), salt meadow, 11.v.2012, 3 ♀♀, P. Kment lgt. et det. (NMPC); Èenkov, Èenkovská step NNR (47°46′07.6″N 18°31′12.0″E, 8277), Stipa steppe on sands with intermixed Ephedra distachya, 13.v.2012, 1 ♀, P. Kment lgt. et det. (NMPC). Remarks. Euro-Siberian species distributed from Sweden and Portugal to the Far East of Russia, Korea, and Tajikistan (KERZHNER & JOSIFOV 1999, AUKEMA et al. 2013). It lives on Galium (usually G. verum) in xerothermic, nutrient-poor grasslands, especially on sandy substrates (WACHMANN et al. 2004, RABITSCH 2007). In Slovakia, the species was recorded by STEHLÍK & HOBERLANDT (1953) from Okolièná na Ostrove (8173) and Èenkov (8277), by DOBŠÍK (1988) from Plešivecká planina (73–7488) and by DOBŠÍK (1991) from Nitra, Kalvária. Here we provide additional Slovak findings, confirming the other records.

Tupiocoris rhododendri (Dolling, 1972)

Material examined. Bohemia bor.: Láznì Libverda (5157), on Rhododendron sp., 9.vii.2013, 3 ♂♂ 5 ♀♀, K. Hradil lgt. et det. (KHMC, NMPC); Snìžník (5250), 605 m a.s.l., in yellow trap near Rhododendron sp., 7.viii.2011, 1 ♂, E. Kula lgt., K. Hradil det. (KHMC). Remarks. Nearctic species (USA: Ohio, Pennsylvania, Tennessee, West Virginia) (HENRY & WHEELER 1976, 1988) introduced to England, where it was discovered in 1971

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 513 P. K MENT ET AL. and described as a new species, Dicyphus rhododendri (DOLLING 1972). In 2002 it was encountered in the Netherlands (AUKEMA et al. 2005), in 2004 in Germany (Baden- Württemberg, HARMUTH 2004; North Rhine-Westphalia, SCHRAMEYER 2004, HOFFMANN 2010), in 2007 in Belgium (AUKEMA et al. 2007), and recently in Denmark (Skipper in AUKEMA et al. 2013). Predator dwelling on various species of Rhododendron (including Azalea), most frequently of the R. ponticum species-group (DOLLING 1972, HENRY & WHEELER 1976), where it preys on the Illinoia (Masonaphis) lambersi (McGillavry, 1960) and other tiny insects. It overwinters in the egg stage, with the adults occurring from June to the beginning of August (DOLLING 1972; AUKEMA et al. 2005, 2007; HOFFMANN 2010).

Tuponia (Tuponia) macedonica Wagner, 1957 (Fig. 23)

Material examined. Moravia mer., Bøezí (7165) near Mikulov, 11.vii.2009, 1 ♂, K. Hradil lgt. et det., A. Carapezza revid. (KHMC). Remarks. Ponto-Mediterranean species distributed in Bulgaria, Macedonia, Greece, European and Asian Turkey, Cyprus and Iran (KERZHNER & JOSIFOV 1999, AUKEMA et al. 2013); recently found introduced into southern Slovakia (Gbelce, 8177) (HRADIL et al. 2008). It lives on Tamarix spp. (WAGNER 1974); in Slovakia it was collected on Tamarix parviflora together with Tuponia (Chlorotuponia) hippophaes (Fieber, 1861) and Tuponia (Tuponia) elegans (Jakovlev, 1867) (KMENT 2004, HRADIL et al. 2008). New species for the Czech Republic.

REDUVIIDAE Peirates hybridus (Scopoli, 1763)

Material examined. Bohemia centr.: Wran [= Vrané nad Vltavou] (6052), 1897, 1 ♂, Nosalek lgt., P. Kment det. (NMPC); Zavisť (6052), no date, 1 ♀, no collector, P. Kment det. (NMPC). Bohemia or.: Litomyšl (6163), 1879, L. Duda [coll.], P. Kment det. (NMPC). Moravia mer.: Silùvky (6864), ‘U køížku’, field road between xerothermic hedges, 9.viii.2009, 1 L, P. Kment lgt. et det. (PKPC); Hodonín, brickyard (48°52′30″N 17°09′01″E, 7168), 13.v.2010, 1 ♀, J. Dolanský lgt., P. Kment det. (VMPC); Klentnice (7165), 17.x.2003, 1 ♂, J. Dolanský lgt. et det., P. Kment revid. (VMPC). Slovakia occ.: Trenèín (70–7174), no date, 3 ♂♂ 2 ♀♀, Èepelák lgt., P. Kment det. (NMPC); Lúka nad Váhom (7373), 30.iii.2002, 1 ♂, M. Mantiè lgt., P. Kment det. (MMHC). Slovakia centr.: Muráò env., Hrdzavá dolina valley (7286), meadow, 28.v.2004, 1 ♀, M. Mantiè lgt., P. Kment det. (MMHC); Zádiel plateau (7390), v.1978, 1 ♀, V. Šilha lgt., Z. Jindra det. (ZJPC); ditto, 22.ix.1995, 1 ♀, M. Škorpík lgt., Z. Jindra det. (ZJPC). Slovakia or.: Kaluža (7198), 25.v.2003, 1 ♀, B. Zbuzek lgt., Z. Jindra det. (ZJPC); Hrhov (7390), 7.x.1994, J. Vávra lgt., P. Kment det. (MMHC); Sirník (7496), 14.ix.1951, 1 ♀, Maøan lgt., P. Kment det. (NMPC); Streda nad Bodrogom, Tarbucka sands (7696), 28.ix.1991, J. Král lgt., P. Kment det. (JVPC). Slovakia mer.: Štúrovo, Hegyfarok [= Belianské kopce hills] (8177), 5.–14.ix.1995, 2 ♂♂ 3 ♀♀ (JVPC, NMPC, PKBC); Kamenica nad Hronom (8178), 16.iii.1997, 1 ♂, Jurèíèek lgt., Z. Jindra det. (ZJPC); Štúrovo, Parkán (81–8278), 23.v.1935, no collector, 1 ♀, P. Kment det. (NMPC); Štúrovo (81–8278), 15.v.1970, 1 ♀, Z. Jindra lgt. et det. (ZJPC). Remarks. Holomediterranean–Central Asiatic species distributed from France and Morocco to Kyrgyzstan, Afghanistan and north-western China (Xinjiang), reaching as far

514 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V north as southern Germany, Czech Republic, and Slovakia (PUTSHKOV & PUTSHKOV 1996, HOFFMANN & MELBER 2003, WACHMANN et al. 2006, LIU et al. 2011, AUKEMA et al. 2013). In central Europe it lives in xerothermic, stony, open habitats, mostly on limestones; in more eastern areas the species seems more hygrophilous, found on the banks of various water reservoirs and streams. It is epigeic, preying largely on other Heteroptera (PUTSHKOV 1987, STEHLÍK & VAVØÍNOVÁ 1997a, WACHMANN et al. 2006, RABITSCH 2007, PUTSHKOV & MOULET 2010). In the Czech Republic, P. hybridus is apparently extinct in Bohemia, not recorded since 1897: Závist u Prahy, environs of Litomyšl (DUDA 1884, 1885; here revised), and Vrané nad Vltavou (NICKERL 1905; here revised). In Moravia, the species is known in the southern, Pannonian region; there are several old records (most recent from 1948) summarized by STEHLÍK & VAVØÍNOVÁ (1997a) and one additional record from 1998 (Ratíškovice – Rudník, 7069) (KMENT & BRYJA 2001). In Slovakia, the species is known from about 20 localities throughout the warm regions of the country, with a gap between records in 1979 and 1993 (DOBŠÍK 1979, STEHLÍK & VAVØÍNOVÁ 1998, ŠTEPANOVIÈOVÁ 2001). In the Czech Republic, the species is classified as “critically endangered” (KMENT & VILÍMOVÁ 2006). Here we provide new recent records from both Moravia and Slovakia.

ARADIDAE Aradus (Aradus) bimaculatus Reuter, 1872 (Fig. 24) Material examined. Bohemia centr.: Praha – Smíchov (5952a), city park, under bark of stump of Aesculus sp., 26.iii.1985, 1 ♀, J. Rùžièka lgt., P. Kment det. (NMPC); Praha – Hostivaø (5953), 31.i.1976, 1 ♂ 4 ♀♀, Z. Kaèenka lgt., Z. Jindra det. (ZJPC). Slovakia mer.: Hajnáèka env., Ragáè hill (7785), 24.viii.2011, 1 ♀, O. Konvièka lgt., P. Kment det. (OKZC). Remarks. Euro-Siberian species (Andorra, Austria, Czech Republic, Finland, France, Germany, Hungary, Poland, Spain, Sweden, Ukraine, and Russia (European Territory, East Siberia)), extending to central Asia (Asian Kazakhstan, Uzbekistan) (HEISS 2001, KANYUKOVA & VINOKUROV 2008, AUKEMA et al. 2013). Mycophageous, the species is associated with deciduous trees (Acer, Alnus, Malus, Quercus, Populus tremula, Tilia), although it has also been recorded from Picea abies (ROUBAL 1958, OSSIANNILSSON 1967, COULIANOS 1989, HEISS & PÉRICART 2007, WACHMANN et al. 2007, ESSER 2010). COULIANOS (1989) reported the species from Populus tremula infested with Entoleuca (= Hypoxylon) mammatum (Ascomycetes: Xylariaceae) in Sweden. ESSER (2010) found specimens of A. bimaculatus collected on the mazegill Daedalea quercina (Basidiomycetes: Fomitopsidaceae), sitting between the gills of the sporocarp and in the gap between the sporocarp and the bark of Quercus robur in Brandenburg. In the Czech Republic, A. bimaculatus was found in 1955–1956 on and Tilia in two localities in central Bohemia, Roztoky u Prahy (5852) and Veltrusy (5751–52) (ROUBAL 1958). KMENT & VILÍMOVÁ (2006) classified the species as “endangered” in the Czech Republic. Here we provide additional records from Bohemia and a first record for Slovakia.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 515 P. K MENT ET AL.

Aradus (Aradus) crenaticollis R. Sahlberg, 1848 (Fig. 25)

Material examined. Bohemia centr.: Milovice (5755), under bark of spruce stumps, 15.iv.2013, 3 ♂♂ 1 ♀, V. Hanzlík lgt., P. Kment det. (VHNC, MNPC). Remarks. Euro-Siberian species known in Austria, Czech Republic, Finland, France, Germany, Italy, Latvia, Norway, Spain, Sweden, Switzerland, Russia (Central and Northern European Territory, Siberia, Far East), Asian Kazakhstan, and Mongolia (HEISS 2001, KANYUKOVA & VINOKUROV 2008, AUKEMA et al. 2013). It prefers coniferous trees, and is known from Picea abies and Pinus sylvestris (SEIDENSTÜCKER 1961, WAGNER 1966, RAMPAZZI & DETHIER 1997, HEISS & PÉRICART 2007). In the Alps it occurs largely in the coniferous zone at medium altitudes on Picea abies (WACHMANN et al. 2007). However, HELIÖVAARA & VÄISÄINEN (1983) noted it from both deciduous and coniferous trees in burned forests. Aradus crenaticollis prefers charred trees after and it is attracted to the small spot fires (e.g. burning ants’ nests) that may continue for several days after passing of a fire front. This attraction is associated with a high migration capacity (COULIANOS 1989; AHNLUND & LINDHE 1992; WIKARS 1992, 1997). WIKARS (1992) reported observations of this species in the immediate vicinity of glowing ants’ nests, sometimes copulating. It probably feeds largely on pyrophilous fungi. However, it has also been found on tree-trunks in sawmills (COULIANOS 1989). Despite its reported pyrophilous tendencies, no photomechanical infrared sensilla, known in some other pyrophilous Aradus spp., have been found in A. crenaticollis (SCHMITZ et al. 2010). In the Czech Republic, the species was previously known from the Šumava Mts. in southern Bohemia, localities Vimperk – Veselka, 3 km N of Boubín (6948) and Kubova Huť, Šeravský les Forest (7048). It was classified as “endangered” in the Red List of the Czech fauna (KMENT & VILÍMOVÁ 2006). Here we provide an additional record, the first for central Bohemia.

Aradus (Aradus) kuthyi Horváth, 1899

Material examined. Bohemia centr.: Vonoklasy (6051), 25.v.2005, 1 ♀, B. Zbuzek lgt., Z. Jindra det. (ZJPC); Praha – Toèná, Šance, Bøežanský dùl valley, 16.iv.1979, 1 ♀, J. Strejèek lgt., Z. Jindra det. (ZJPC); Praha – Toèná, Šance (6052a), 9.vi.2001, 1 ♂, J. Strejèek lgt., P. Kment det. (OBPC). Moravia mer.: Senorady (1.5 km N), Oslava river valley, Levnov castle ruin (6863c), rocks, viii.2012, 1 ♀, V. Kubáò lgt., P. Kment det. (NMPC); Podyjí NP, Podmolí env., Lišèí skála rocks (7161), 3.vii.1995, 1 ♀, S. Bílý lgt., P. Kment det. (NMPC). Slovakia mer.: Plášťovce (7879), 30.v.1993, 1 ♀, B. Zbuzek lgt., Z. Jindra det. (ZJPC). Remarks. Balkan-Pannonian species, known only from Albania, Austria, Bosnia and Herzegovina, Czech Republic, Greece, Hungary, and Slovakia (HEISS 2001, AUKEMA et al. 2013). It occurs on Juglans regia, Quercus spp. (Q. cerris, Q. petrea, Q. robur), and Pinus sylvestris (HOBERLANDT 1956, ŠTYS 1976, ADLBAUER 1992, HEISS & PÉRICART 2007, WACHMANN et al. 2007). In the Czech Republic, it is recorded only from central Bohemia, Závist near Praha (6052), where it was found under a fine scale of bark on a thin Pinus sylvestris twig, the tree isolated on a xerothermic, rocky slope (ŠTYS 1976). In Slovakia, A. kuthyi was collected in the Kováèovské kopce Hills near Štúrovo (8177) (HOBERLANDT 1956, STEHLÍK & HEISS 2001) and Pstruša (7481) (STEHLÍK & HEISS 2001).

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It is regarded an endangered species in both the Czech Republic (KMENT & VILÍMOVÁ 2006) and Slovakia (ŠTEPANOVIÈOVÁ & BIANCHI 2001). New species for Moravia, and second record for Bohemia.

Aradus (Aradus) mirus Bergroth, 1894 Material examined. Bohemia bor.: Julèín, Na Èernèí NR (5452), Pulsatillo-Pinetea, 1.vi.1999, 1 L, I. Malenovský lgt., P. Kment det. (NMPC). Bohemia occ.: Skláøe near Mariánské Láznì (6042), 20.v.2001, 1 ♀, J. Strejèek lgt., P. Kment det. (OBPC); Obora near Plzeò (6146), 1976, 1 ♀, no collector, L. Hoberlandt det. (NMPC). Bohemia centr.: Køivoklát, Baba hill [= Na Babì NR] (5949), 28.v.1988, 1 ♀, S. Bílý & J. Jelínek lgt., P. Kment det. (NMPC); Sedlèany env., Kounova [= Kosova] Hora (6352), 27.v.2005, 1 ♀, J. Strejèek lgt., P. Kment det. (NMPC); Louòovice pod Blaníkem, Velký Blaník hill (6355), 600 m a.s.l., 2.vii.1988, 1 ♀, J. Strejèek lgt., Z. Jindra det. (ZJPC); Sedlice, Želivka dam, Sedlické údolí valley (6356), 28.v.1966, 1 ♀, J. Strejèek lgt., P. Kment det. (NMPC). Bohemia mer.: Vráž near Písek (6650), 400 m a.s.l., 19.–25.vi.2010, 1 ♀, M. Barták lgt., Z. Jindra det. (ZJPC). Moravia occ.: Kost. Mysletice [= Kostelní Myslová] (6858), 8.–13.x.1973, Z. Šustek lgt., P. Kment det. (MMBC: coll. Dobšík). Moravia mer.: Bzenec – Pøívoz (7069), beating branches of Pinus, 3.iv.2010, 1 ♀, J. Ch. Vávra lgt., P. Kment det. (OMOC). Slovakia occ.: Malacky (7568), 28.xi.1987, 1 ♀, I. Jeniš lgt., P. Kment det. (PKBC). Remarks. Species endemic to the Pannonian lowlands (Austria, Moravia, Hungary, Slovakia), extending into Bohemia (HEISS 2001). It lives on pines (Pinus sylvestris, P. nigra) on xerothermic localities in lowlands. Unlike most other bark bugs, A. mirus very probably sucks direct from the phloem of its host plants. Both adults and larvae may overwinter and may be found throughout the year (HEISS & PÉRICART 2007, RABITSCH 2007, WACHMANN et al. 2007). The species is considered endangered in the Czech Republic (KMENT & VILÍMOVÁ 2006) as well as in Lower Austria (RABITSCH 2007). In Bohemia, it was previously known from Libøice near Vrané nad Vltavou (HOBERLANDT 1944a, c, 1956) and Svádov (5350) (HOBERLANDT 1956), in Moravia from Mohelno (Mohelenská hadcová step NNR, 6863) (STEHLÍK 1946, STEHLÍK & HEISS 2000) and Nemotice (Vysoká Hill) (STEHLÍK & HEISS 2000), and in Slovakia from Trenèín (70–7174) and Košice (72–7393) (HOBERLANDT 1944b, c). Here we provide recent records from all three Lands.

Aradus (Aradus) signaticornis R. Sahlberg, 1848 Material examined. Bohemia centr.: Praha – Bøevnov, Pod Královkou (5952), at window in a house, 5.v.2003, 1 ♀, J. Rùžièka lgt., P. Kment det. (PKBC). Remarks. Holarctic species, distributed from Sweden and France to the Far East of Russia, Korea, and south-western China (Yunnan) (HEISS 2001, 2010; KANYUKOVA & VINOKUROV 2008; AUKEMA et al. 2013). It is a pioneer species, occupying habitats in early phases of succession, such as industrial wastelands, spoil heaps after coal mining, forest clearings, and forest fire sites (PERSSON 1966, ŠTUSÁK 1976, HELIÖVAARA & VÄISÄINEN 1983, BALS et al. 1997, WIKARS 1997, ABS et al. 1999, HJÄLTÉN et al. 2006, HEISS & PÉRICART 2007, WACHMANN et al. 2007). It has been recorded on both coniferous and deciduous trees (Pinus, Larix, Betula) (PERSSON 1966, BALS et al. 1997, WIKARS

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1997, HEISS & PÉRICART 2007, WACHMANN et al. 2007). In the Czech Republic, the species was previously known only from Bohemia, locality Zbùch (6345) 13 km south- west of Plzeò; this record was based on a single specimen bred from a 5th-instar larva found under a stone in a large heap of stones carted away from the coal mine, the locality being almost devoid of vegetation, with only scattered young birches (Betula sp.) (ŠTUSÁK 1976). Here we provide the second record for the Czech Republic.

LYGAEIDAE Arocatus melanocephalus (Fabricius, 1798) (Fig. 26) Material examined. Slovakia mer.: Zlatná na Ostrove – Ve¾ký Lel, Ve¾ký Lel Island (N 47°44′51.6″ E 17°56′33.8″, 8273), Zlatòanský luh floodplain forest, beating lower branches of Ulmus laevis, 12.v.2012, 1 ♀, E. Líznarová lgt., P. Kment det. (NMPC). Remarks. Northern Mediterranean species distributed from Germany and Portugal to Iran, recently discovered in Xinjiang (NW China) as well (PÉRICART 2001, AUKEMA et al. 2013, GAO et al. 2013). It develops on Ulmus, both adults and larvae sucking nourishment from seeds. It overwinters in the adult stage, seeking shelter under the loose bark of trees (, oaks, etc.), often in aggregations, or in crevices in tree-trunks, in pine- cones, or in layers of dead leaves (PUTSHKOV 1969, PÉRICART 1999a). In central Europe it has also been recorded from Quercus and Alnus (WACHMANN et al. 2007). In the first decade of the 21st century, sudden mass occurrences and subsequent intrusions into human dwellings by A. melanocephalus, previously considered a very rare species, were observed during summer in urban environments in northern Italy (REGGIANI et al. 2005, MAISTRELLO et al. 2006, DUTTO & CARAPEZZA 2011) and Germany (HOFFMANN & TERME 2012). There appears to be a link between this phenomenon and recent climate warming, which seems to favour the survival of overwintering adults during winter and developing larval stages in spring but endanger both larvae and adults (intolerant to temperature >30°C) on hot summer days and force them to seek cool aestivation shelters inside buildings (MAISTRELLO et al. 2006). On the other hand, DUTTO & CARAPEZZA (2011) hypothesised that the migration into houses is induced by mean temperatures similar to those in autumn, which apparently causes the insects to seek out places suitable for hibernation. There is only one record of this species for Slovakia, a single specimen collected in the Svätý Peter oak forest (S Slovakia, 8175d) on 1.vii.1959 (ŠTEPANOVIÈOVÁ 2003). Here we confirm the occurrence of the species in Slovakia after an interval of 53 years.

Melanocoryphus albomaculatus (Goeze, 1778)

Material examined. Moravia mer.: Moravské Bránice (6964), 6.ix.2008, 1 ♀, K.Hradil lgt. et det. (KHMC); Hnanice, Šobes (7162), 1.v.2004, 1 ♂, J. Vilímová lgt., P. Kment det. (NMPC). Remarks. Northern Mediterranean–Central Asiatic species distributed from the Netherlands, Portugal, and Morocco in the west to Iran and central Asia in the east (PÉRICART 2001). Epigeic, thermophilous species, living chiefly on sunlit rocky hillsides

518 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V and in clearings, where it is found under lichens, moss, dry leaves, stones, etc. It feeds largely on Vincetoxicum hirundinaria (especially the larvae), although other plants (e.g., Digitalis, Senecio) may serve as well, feeding mostly on seeds (STEHLÍK & VAVØÍNOVÁ 1996, 1997b; PÉRICART 1999a; WACHMANN et al. 2007). In Moravia, there was a long gap in records between 1962 and 1992 and in the 1990s the species was documented only in the Podyjí National Park (STEHLÍK & VAVØÍNOVÁ 1997b, KMENT et al. 2003). Only recently did MALENOVSKÝ et al. (2011) record M. alboacuminatus from Budkovické skály Rocks (6964) near Budkovice outside Podyjí NP. In the Czech Republic, the species is classified as “critically endangered” (KMENT & VILÍMOVÁ 2006). Here we provide two additional records.

Melanocoryphus tristrami (Douglas et Scott, 1868) (Fig. 27) Material examined. Slovakia mer.: Èenkov, Èenkovská step NNR (47°46′07.6″N, 18°31′12.0″E, 8277), Stipa steppe on sands, 13.v.2011, 1 ♀ (ma), P. Kment lgt. et det. (NMPC). Remarks. Ponto-Mediterranean–Central Asiatic species distributed from Hungary and Croatia to Iran and central Asia (PÉRICART 2001, AUKEMA et al. 2013). It is a xerophilous, epigeic species dwelling among plant detritus, feeding on the seeds of various plants (PUTSHKOV 1969, PÉRICART 1999b). In Slovakia, the species was previously known from only a single record from Belianské kopce Hills (= Hegyfarok) near Štúrovo (8177) (KMENT et al. 2003). Here we provide a second Slovak locality.

GEOCORIDAE Geocoris (Piocoris) erythrocephalus (Lepeletier et Serville, 1825) (Fig. 28) Material examined. Moravia mer.: Poštorná, environs of Boøí les railway station (48°44′09.98″N 16°52′10.63″E, 7267), sweeping of xerothermic ruderal vegetation on slopes around sludge bed, 18.viii.2013, 1 ♂, P. Kment lgt. et det. (NMPC). Remarks. Holomediterranean species extending to the southern areas of central Europe (Austria, Hungary, Slovakia), Iran and Kazakhstan (PÉRICART 2001, AUKEMA et al. 2013). Predatory species, largely epigeic, overwintering in the adult stage (STEHLÍK & VAVØÍNOVÁ 1996, PÉRICART 1999a, WACHMANN et al. 2007). In the Mediterranean area it is quite an euryecious species, also found in humid habitats (PÉRICART 1999a, WACHMANN et al. 2007), while in central Europe it appears bound to xerothermic habitats with scattered groups of vegetation, such as disused vineyards and pastures, and steppe vegetation on the slopes of readily-warmed rocks (especially andesite) (STEHLÍK & VAVØÍNOVÁ 1996, ŠTEPANOVIÈOVÁ 2001, ŠTEPANOVIÈOVÁ & BIANCHI 2003, RABITSCH 2007). In Austria, the species is known from a few localities in Burgenland, Styria (ADLBAUER 1995, 1997), and Lower Austria, where it was first recorded in 2003 in an abandoned sand-pit near Lichtenegg (RABITSCH 2003, 2007). In Slovakia, the species is known from a number of localities, although all of them are situated in southern parts of the country (STEHLÍK & VAVØÍNOVÁ 1996, ŠTEPANOVIÈOVÁ 2001, RUS & KMENT 2007). The westernmost Slovak localities are the environs of Bratislava (7868–69) (ORTVAY

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1902, as Pozsóny) and Jurský Šúr near Svätý Jur (7769) (ORSZÁGH 1966), both about 60 km south of Bøeclav and Kòazí vrch Hill (7373) in the Tematínské kopce hills (ŠTEPANOVIÈOVÁ 2001) 77 km east of Bøeclav. The species was listed for Moravia by HOBERLANDT (1977a) based on ROUBAL (1964, 1968), but its occurrence in Moravia was rejected by STEHLÍK & VAVØÍNOVÁ (1997b). Geocoris erythrocephalus might be considered among the species recently and naturally expanding their distribution ranges in central Europe due to changes in climatic conditions that favour them (see e.g. RABITSCH 2008). New species for the Czech Republic.

OXYCARENIDAE Tropidophlebia costalis (Herrich-Schaeffer, 1850)

Material examined. Bohemia bor.: Kleneè, Kleneè NNM (5651), 17.v.2008, 1 ♀, K. Hradil lgt. et det. (KHMC). Bohemia mer.: Lužnice env., 0.5 km NE of the village, Slepièí vršek hill NM (6954), sandy dune, under Calluna, 2.v.2008, 1 ♀, Z. Kejval lgt., P. Baòaø det. (MCHD). Moravia mer.: Bzenec – Pøívoz, Vojenské cvièištì NM (48°57′30″N 17°17′25″E, 7069), sands in former military training area, 15.v.2010, 1 ♂, P. Kment lgt et det. (NMPC); Havraníky, Havranické vøesovištì heathland (7162), 4.vii.1995, pitfall trap, 3 ♂♂ 3 ♀♀ (one couple in copula), 16.viii.1995, 2 ♀♀, pitfall trap, A. Reiter lgt., P. Kment det. (NMPC). Remarks. Euro-Siberian species distributed from Sweden and France to East Siberia and Mongolia (PÉRICART 2001, AUKEMA et al. 2013). It dwells on aeolian sands, sterile soils and sunlit rocks with sparse vegetation, lichens and plant litter, feeding on the seeds of various plants (STEHLÍK & VAVØÍNOVÁ 1996, 1997b; PÉRICART 1999b, WACHMANN et al. 2007). Its distribution in Moravia is limited to the warmest southern parts of the territory (STEHLÍK & VAVØÍNOVÁ 1997b, KMENT et al. 2003). However, in Bohemia the species was previously known from only a single locality, Kleneè NNM (5651) in the Labe (= Elbe) lowlands. In the Czech Republic the species is classified as “near-threatened” (KMENT & VILÍMOVÁ 2006). Here we confirm its occurrence in Kleneè and provide an additional record for southern Bohemia.

COREIDAE Coriomeris scabricornis scabricornis (Panzer, 1805)

Material examined. Bohemia bor.: Kleneè, Kleneè NNM (5651), 11.vii.2008, 1 ♂, K. Hradil lgt. et det. (KHMC). Bohemia centr.: Stará Boleslav (5854), former military training area on sands, 17.ix.1995, 3 ♂♂ 4 ♀♀, P. Kapitola lgt., K. Hradil det. (KHMC). Moravia mer.: Mutìnice env. (70–7168), 9.viii.1975, 1 m, J. Adámek lgt., Z. Jindra det. (ZJPC); Bzenec, Vojenské cvièištì NM (48°57′30″N 17°17′25″E, 7069), pitfall trap, 12.vi.2009, 1 ♂, 11.vii.2009, 1 ♂, P. Chlapek & O. Èížek lgt., K. Hradil det. (KHMC); Rohatec – Kolonie (7169), 7.vi.1985, 1 ♀, Z. Jindra lgt. et det. (ZJPC); Lednice env. (71–7266), 3.v.1975, 1 ♀, Z. Jindra lgt. et det. (ZJPC). Slovakia occ.: Mikulášov near Lakšárska Nová Ves, 9.vii.1991, 2 ♂♂, P. Kapitola lgt., K. Hradil det. (KHMC). Slovakia or.: Hrhov env. (7390), 25.vi.1976, 1 ♂, R. Pucholt lgt. (ZJPC). Remarks. Holopalaearctic species distributed from Sweden and Portugal to Korea and Japan, absent from North Africa (DOLLING 2006, AUKEMA et al. 2013). It lives on Fabaceae (e.g. Trifolium and Medicago), Lamiaceae (Thymus, Ajuga) and Asteraceae (Artemisia, Helichrysum) (STEHLÍK 1988, WACHMANN et al. 2007), mainly on aeolian sands in Moravia, but has also been found in xerothermic habitats on rocky substrates,

520 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V largely restricted to the Pannonicum (STEHLÍK 1988). Coriomeris scabricornis used to be quite a common species in suitable localities in southern Moravia until the 1970s, but disappeared in the 1980s and has not been found in Moravia since 1986 (STEHLÍK 1988). Some recent records for Bohemia were published by RUS (2005) and KMENT & BAÒAØ (2012). There is only a single recent record from Lower Austria as well (RABITSCH 2002, 2007). The species was rediscovered in Moravia only recently, in 2009 on aeolian sands in Bzenec (this paper) and in 2010 on natural vegetation in the Kamenárka limestone quarry near Štramberk (6474) in north-eastern Moravia (LIS 2012). In the Czech Republic the species is classified as “critically endangered” (KMENT & VILÍMOVÁ 2006).

ALYDIDAE Camptopus lateralis (Germar, 1817) (Fig. 29) Material examined. Moravia mer.: Poštorná, environs of Boøí les railway station (48°44′09.98″N 16°52′10.63″E, 7267), sweeping of xerothermic ruderal vegetation on slopes around sludge bed, 18.viii.2013, 2 ♀♀, P. Kment lgt. et det. (MMBC, NMPC). Remarks. A West-Palaearctic species, distributed from France, Portugal, Morocco and Macaronesia in the west to Northwest China, Afghanistan, Pakistan and India in the east (DOLLING 2006, AUKEMA et al. 2013). In central Europe the species is known from Switzerland, Liechtenstein, Austria, Slovakia, and Hungary, with old and dubious records from Belgium and Germany (HOFFMANN & MELBER 2003, DOLLING 2006, AUKEMA et al. 2013). In the Mediterranean area it is quite a euryecious species, recorded from plants of various families, although it shows a certain preference for Fabaceae (MOULET 1995). In central Europe the species is bound to xerothermic habitats with scattered groups of vegetation – sands, salt meadows, and especially slopes of readily-warmed rocks (limestone, andesite, basalt) covered with steppe vegetation. It has been recorded on various plants, although predominantly on Fabaceae (Lotus, Coronilla, Medicago, Trifolium, Onobrychis, etc.) (STEHLÍK & VAVØÍNOVÁ 1995, RABITSCH 2007, WACHMANN et al. 2007). It overwinters in the adult stage, with only one generation per year in central Europe, while more generations are produced in more southern regions of its range (MOULET 1995, STEHLÍK & VAVØÍNOVÁ 1995, RABITSCH 2007). In Austria the species is known from Burgenland and Lower Austria, although it was considered extinct in Lower Austria by RABITSCH (2007), but a single specimen was re-discovered in 2012 (W. Rabitsch, pers. comm.). In Slovakia, C. lateralis is known from a number of localities in the south-west of the country and there is a solitary record from the south-east (STEHLÍK & VAVØÍNOVÁ 1995, ŠTEPANOVIÈOVÁ 2001, ŠTEPANOVIÈOVÁ & BIANCHI 2003, RUS 2005). The westernmost Slovak localities are Devínská Kobyla Hill in Bratislava (7867–68) (BALTHASAR 1942), about 61 km south of Bøeclav and Kòazí vrch Hill (7373) in the Tematínské kopce hills (ŠTEPANOVIÈOVÁ 2001), situated 77 km east of Bøeclav. The species was listed for Moravia by HOBERLANDT (1977a) in error (see STEHLÍK 1988). Camptopus lateralis may also be considered among the species recently and naturally expanding their distribution ranges in central Europe due to changes in climatic conditions that favour them (see e.g. RABITSCH 2008). New species for the Czech Republic. Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 521 P. K MENT ET AL.

PENTATOMIDAE Aelia rostrata Boheman, 1852

Material examined. Moravia mer.: Klentnice, Tabulová hora NNR (7165), W slope, 18.v.2004, 1 ♂, O. Balvín lgt. et det., P. Kment revid. (OBPC); Sedlec, Skalky u Sedlece NM (48°46′28.30″N 16°40′21.59″E, 7266), steppe, 1.v.2013, 1 ♂ 1 ♀, M. Horsák, P. Kment & Biotým lgt. (MHBC, NMPC); Poštorná, environs of Boøí les railway station (N 48°44′09.98″ E 16°52′10.63″, 7267), sweeping of xerothermic ruderal vegetation on slopes around sludge bed, 18.viii.2013, 1 ♂, P. Kment lgt. et det. (NMPC). Slovakia mer.: Nitra, Lupka NR (7674d), 11.iv.2008, 1 ♂, J. Cunev lgt., P. Kment det. (JCNS); Nitra, Kalvária hill (7774a), 20.ix.2010, 1 ♀, J. Cunev lgt., P. Kment det. (JCNS); Nesvady, Líšèie diery NR (8074d), 30.viii.2008, 1 ♀, J. Cunev lgt., P. Kment det. (JCNS); Martovce env. (8174a), 23.viii.2010, 1 ♀, J. Cunev lgt., P. Kment det. (JCNS); Kováèov (8178), 16.v.1953, 1 ♀, J. Štaif lgt., Z. Jindra det. (ZJPC); Salka, Sovie Vinohrady (8178), steppe, sweeping, 10.ix.2001, 1 ♂, M. Mantiè lgt., P. Kment det. (MMHC); Èenkov, Èenkovská step NNR (47°46′07.6″N 18°31′12.0″E, 8277), Stipa steppe on sands, 13.v.2011, 6 ♂♂, P. Kment lgt. et det. (MHBC, NMPC); Mužla env., Jurský Chlm (8277), steppe, sweeping, 10.ix.2000, 1 ♀, M. Mantiè lgt., P. Kment det. (MMHC). Slovakia or.: Ladmovce (7596), 7.v.2000, 1 ♂ 1 ♀, V. Zeman lgt., P. Kment det. (VZPC). Remarks. A widely distributed West-Palaearctic species with the centre of its distribution in the Mediterranean, but reaching as far as Denmark, Sweden, and Byelorussia in the north (RIDER 2006, AUKEMA et al. 2013). It prefers xerothermic grasslands, feeding on Poaceae (STEHLÍK 1985, DERJANSCHI & PÉRICART 2006, WACHMANN et al. 2008). A remarkable decline in the distribution of A. rostrata was observed in Germany and Austria since the middle of 20th century, with the species becoming extinct in Germany and recent records from just eastern Austria (GÜNTHER et al. 1998, RABITSCH 2007, WACHMANN et al. 2008). The same phenomenon occurred in the Czech Republic, well documented by STEHLÍK (1985) in Moravia, where the species was still widespread but quite rare in the 1940–1950s, but had not been previously found since 1963. This led to the species′ being considered “regionally extinct” in the Czech Republic (KMENT & VILÍMOVÁ 2006). In Slovakia, it became very rare in the 1970–1980s, although it seems to have maintained a continuous presence there, with an increasing number of records since 1990 (ŠTEPANOVIÈOVÁ 1977, 1991; STEHLÍK & VAVØÍNOVÁ 1993). Here we provide the first recent records of the species for the Czech Republic after an interval of 41 years.

Eurydema fieberi Fieber, 1837 (Fig. 30) Material examined. Bohemia bor.: Ústí nad Labem – Brná, Sluneèní stráò NM (5350), rocky steppe, 22.vi.2011, 2 ♂♂, J. Moravec lgt., P. Kment det. (NMPC); Lovosice, Lovoš NNR (5450), rocky steppe under the top, 22.iv.2011, 1 spec., photographed by J. Vítek, P. Kment det. Bohemia centr.: Vrané nad Vltavou, Zvolská homole NR (6052), 28.iii.1998, 1 ♂, M. Øezáè lgt., P. Kment det. (VMPC). Moravia mer.: Mohelno, Mohelenská hadcová step NPR (49°06′31.53″N 16°11′12.99″E, 6863), on and under Alyssum montanum, 11.v.2013, 11 spec. observed, M. Horsák, P. Kment & Biotým lgt. (1 ♂ in MHBC, 2 ♂♂ 1 ♀ in MVBC, 2 ♂♂ in NMPC); Klentnice, Rùžový vrch NNR (7165), 12.vii.2010, 1 ♂, K. Hradil lgt. et det. (KHMC). Slovakia mer.: Tríbeè Mts., Štitáre env., Žibrica (7674–75), 1.viii.1982, 1 ♂, J. Cunev lgt., P. Kment det. (JCNS); Plášťovce (7879), 30.v.1990, 1 ♂, P. Kapitola lgt., K. Hradil det. (KHMC). Remarks. Holomediterranean–Central Asiatic species distributed from Portugal and Morocco to the Northwest Territory of China, Afghanistan, and India (RIDER 2006, AUKEMA et al. 2013). Eurydema fieberi lives on plants of the family Brassicaceae, e.g. Erysimum spp., Biscutella laevigata, Leucosinapis (= Sinapis) alba, Isatis tinctoria,

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Erophila (= Draba) verna, Aurinia (= Alyssum) saxatilis, Cardaria draba, Sisymbrium spp., Lepidium spp., Thlaspi spp., and Barbarea vulgaris subsp. arcuata (STEHLÍK 1986, STEHLÍK & VAVØÍNOVÁ 1994, DERJANSCHI & PÉRICART 2006, WACHMANN et al. 2008). Here we report it for the first time on Alyssum montanum. KMENT & VILÍMOVÁ (2006) considered the species “regionally extinct” in the Czech Republic, based on STEHLÍK (1986). However, it was subsequently demonstrated that the species survives in the Èeské Støedohoøí Mts. and the River Vltava valley in the environs of Prague in Bohemia (KMENT & JINDRA 2008). Despite an intensive sampling effort, the species has not been recorded in Moravia since 1952 (records from Pavlovské vrchy Hills), in the Mohelno locality even since 1942 (STEHLÍK 1986). Here we provide a recent Moravian record after an interval of 61 years.

Errata to previous paper KMENT & BAÒAØ (2012): The following changes are required: • p. 519. Cimex pipistrelli Jenyns, 1839. Add a published record from Luhaèovice (6872) for Myotis myotis (BARTONIÈKA 2010). Overlooked.

• p. 522. Aneurus (Aneurus) laevis (Fabricius, 1775). The record from Starý Hrozenkov, Skalka quarry, is based on a misidentification of Aneurus (Aneurodes) avenius (Dufour, 1833). There is no record of A. laevis in the Bílé Karpaty Protected Landscape Area; the number of species occurring in the Bílé Karpaty PLA is therefore reduced to 500.

Acknowledgements We are indebted to the following colleagues who donated their material or made their collections available for our studies: Ludvík Bobot (Zlín), Jan Dolanský (Museum of Eastern Bohemia in Pardubice), Libor Dvoøák (Municipal Museum, Mariánské Láznì), Václav Hanzlík (Neratovice), Michal Horsák (Department of Botany and Zoology, Masaryk University, Brno), Marion Mantiè (Hluèín), Petr Komzák (Brno), Ondøej Konvièka (Zlín), Martin Vašíèek (Brno), Jiøí Ch. Vávra (Museum of Ostrava, Ostrava), and Vladimír Zeman (Pøerov). We also extend our thanks to Vít Grulich (Department of Botany and Zoology, Masaryk University, Brno) for information on the distribution of certain host plants, and Berend Aukema (NCB Naturalis, Leiden, the Netherlands), Gavril Marius Berchi (West University of Timiºoara, Timiºoara, Romania), Pál Boda (Trans Tisza Region Environmental, Nature Protection and Water Inspectorate, Debrecen, Hungary), Aleksander Herczek (Silesian University, Katowice, Poland), Elöd Kondorosy (Georgikon Faculty, University of Pannonia, Keszthely, Hungary), Wolfgang Rabitsch (Umweltbundesamt, Abt. Biologische Vielfalt & Naturschutz, Vienna, Austria), Pavel Štys and Jitka Vilímová (both Charles University, Prague, Czech Republic) for valuable information and/or providing inaccessible reprints. This study was partly supported by grants from the Ministry of Culture of the Czech Republic, ref. DKRVO MK-S 760/2013/11 OVV, to the National Museum, Prague (P. Kment) and ref. MK000094862 to the Moravian Museum, Brno (P. Baòaø).

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Figs 10–17. Habitus of some Heteroptera recorded in the Czech Republic and Slovakia. 10 – Arctocorisa germari germari (Fieber, 1848), ♂, Pøebuz (MMLC) (8.7 mm); 11 – Microvelia buenoi Drake, 1920, ♂ (apterous), Hrabanovská èernava NNR (2.0 mm); 12 – M. buenoi, ♀ (apterous), Kokoøínský dùl NR (1.7 mm); 13 – M. buenoi, ♂ (macropterous), Hrabanovská èernava NNR (2.1 mm); 14 – M. buenoi, ♀ (macropterous), Hrabanovská èernava NNR (1.9 mm); 15 – M. pygmaea (Dufour, 1833), ♂ (apterous), Most pri Bratislave (MSHC); 16 – M. pygmaea, ♀ (apterous), Ivaò, Nové Mlýny (NMPC); 17 – M. pygmaea, ♀ (macropterous), Travní Dvùr SAC (MSHC). (Photo: P. Kment.)

524 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

Figs 18–23. Habitus of some Heteroptera recorded in the Czech Republic and Slovakia. 18 – Salda henschi (Reuter, 1891), ♂, Rejvíz (NMPC) (4.7 mm); 19 – Agramma atricapillum (Spinola, 1837), ♀, Èiližské mokrade (NMPC) (2.8 mm); 20 – Hyoidea notaticeps Reuter, 1876, ♂, Èenkov (NMPC) (3.8 mm); 21 – Harpocera hellenica Reuter, 1876, ♀, Mohelno (NMPC) (4.4 mm); 22 – Isometopus mirificus Mulsant et Rey, 1879, ♀, Tvrdonice (NMPC) (2.3 mm); 23 – Tuponia macedonica Wagner, 1957, ♂, Bøezí (KHMC) (3.3 mm). (Photo: P. Kment.)

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Figs 24–30. Habitus of some Heteroptera recorded in the Czech Republic and Slovakia. 24 – Aradus bimaculatus Reuter, 1872, ♀, Hajnaèka (OKVC) (5.0 mm); 25 – Aradus crenaticollis R. Sahlberg, 1848, ♀, Milovice (VHNC) (8.2 mm); 26 – Arocatus melanocephalus (Fabricius, 1798), ♂, Ve¾ký Lel (NMPC) (6.9 mm); 27 – Melanocoryphus tristrami (Douglas et Scott, 1868), ♀, Èenkov (NMPC) (5.6 mm); 28 – Geocoris erythrocephalus (Lepeletier et Serville, 1825), ♂, Poštorná (NMPC) (3.3 mm); 29 – Camptopus lateralis (Germar, 1817), ♀, Poštorná (NMPC) (12.7 mm); 30 – Eurydema fieberi Fieber, 1837, ♂, Mohelno (NMPC) (7.0 mm). (Photo: 24, 26–30 P. Kment; 25 V. Hanzlík.)

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References

ABS M., SCHERK A. & ZEIß A. 1999: Bergehalden im Ruhrgebiet – eine Oase für Tiere? Biologie in Unserer Zeit 29: 346–352. ADLBAUER K. 1992: Neue Wanzen für die Steiermark und für Österreich (Heteroptera). (3. Beitrag zur Faunistik steirischer Wanzen). Mitteilungen des Naurwissenschaftlichen Vereins für Steiermark 122: 173–176. ADLBAUER K. 1995: Der Reliktstandort am Steinbruch Klausen bei Bad Gleichenberg – ein neues Naturschutzgebiet. Jahresbericht Landesmuseum Joanneum 24: 45–63. ADLBAUER K. 1997: Neue Wanzen für die Steiermark, das Burgenland und Österreich (Heteroptera). (6. Beitrag zur Faunistik steirischer Wanzen). Mitteilungen des Naturwissenschaftlichen Vereins für Steiermark 127: 157–162. AHNLUND H. & LINDHE A. 1992: Hotade vedinsekter i barrskoglandskapet – några synpunkter utifrån studier av sörmländska brandfält, hällmarker och hyggen. (Endangered wood-living insects in coniferous forests – some thoughts from studies of forest-fire sites, outcrops and clearcutings in the province Sörmland, Sweden). Entomologisk Tidskrift 113: 13–23 (in Swedish, English summary). ANDERSEN N. M. 1995: Infraorder Gerromorpha Popov, 1971 – semiaquatic bugs. Pp. 77–114. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1. The Netherlands Entomological Society, Amsterdam, xxvi + 222 pp. AUKEMA B., BOS F., HERMES D. & ZEINSTRA PH. 2005: Nieuwe en interessante nederlandse wantsen II, met een geactualiseerde naamlijst (Hemiptera: Heteroptera). (New and interesting Dutch bugs II, with an updated checklist (Hemiptera: Heteroptera)). Nederlandse Faunistische Mededelingen 23: 37–76 (in Dutch, English summary). AUKEMA B., BRUERS J. M. & VISKENS G. M. 2007: Nieuwe en zeldzame Belgische wantsen II (Hemiptera: Heteroptera). [New and rare Belgian true bugs (Hemiptera: Heteroptera)]. Bulletin van de Koninklijke Belgische Vereniging voor Entomologie 143: 83–91 (in Dutch, English and French summaries without titles). AUKEMA B., CUPPEN J. G. M., NIESER N. & TEMPELMAN D. 2002: Verspreidingsatlas Nederlandse wantsen (Hemiptera: Heteroptera). Deel I: , Nepomorpha, Gerromorpha & . [Distribution atlas of true bugs of the Netherlands (Hemiptera: Heteroptera). Volume I: Dipsocoromorpha, Nepomorpha, Gerromorpha & Leptopodomorpha]. European Invertebrate Survey – Nederland, Leiden, 169 pp. AUKEMA B. & RIEGER CH. (eds.) 1995: Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1. Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha and Leptopodomorpha. The Netherlands Entomological Society, Amsterdam, xxvi + 222 pp. AUKEMA B. & RIEGER CH. (eds.) 1996: Catalogue of the Heteroptera of the Palaearctic Region. Vol. 2. Cimicomorpha I. The Netherlands Entomological Society, Amsterdam, xiv + 361 pp. AUKEMA B. & RIEGER CH. (eds.) 1999: Catalogue of the Heteroptera of the Palaearctic Region. Vol. 3. Cimicomorpha II. The Netherlands Entomological Society, Amsterdam, xiv + 577 pp. AUKEMA B. & RIEGER CH. (eds.) 2001: Catalogue of the Heteroptera of the Palaearctic Region. Vol. 4. Pentatomomorpha I. The Netherlands Entomological Society, Amsterdam, xiv + 346 pp. AUKEMA B. & RIEGER CH. (eds.) 2006: Catalogue of the Heteroptera of the Palaearctic Region. Vol. 5. Pentatomomorpha II. The Netherlands Entomological Society, Amsterdam, xiii + 550 pp. AUKEMA B., RIEGER CH. & RABITSCH W. 2013: Catalogue of the Heteroptera of the Palaearctic Region. VI. Supplement. The Netherlands Entomological Society, Amsterdam, xxiii + 629 pp. BAKONYI G., CSÖLLE C., FABÓK V., FÖLDESSY M., HUFNAGEL L., KONDOROSY E., RÉDEI D., TÖLGYESINÉ-NELL T., VARGA I. & VÁSÁRHELYI T. 2002: The Heteroptera fauna of the Fertõ-Hanság National Park. Pp. 325–350. In: MAHUNKA S. (ed.): The Fauna of the Fertõ-Hanság National Park I–II. Hungarian Natural History Museum, Budapest, 829 pp. BALS J., BENHOLZ J. & SCHWERK A. 1997: Nachweise von Aradus signaticornis, R. Sahlberg, 1848 in Nordrhein-Westfalen. (Insecta: Heteroptera: Aradidae). Entomologische Mitteilungen aus dem Löbbecke- Museum und Aquazoo 8: 43–44. BALTHASAR V. 1937: Slovenské ploštice. Katalog a pokus o rozbor složek fauny slovenských Heteropter. (Die Heteropteren der Slowakei. Ein katalog der faunistischen Komponenten der Slowakischen Heteropteren). Bratislava 11: 194–249 (in Czech, German summary).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 527 P. K MENT ET AL.

BALTHASAR V. 1942: Pozoruhodné nálezy ploštic na Moravì a na Slovensku. (Über einige beachtenswerte Wanzen-Arten aus Mähren und Slowakei). (Opuscula heteropterologica II). Entomologické Listy (Brno) 5: 25–29 (in Czech, German summary). BALVÍN O. 2007: Faunistic records from the Czech Republic – 231. Heteroptera: Tingidae. Klapalekiana 43: 200. BALVÍN O., ŠEVÈÍK M., JAHELKOVÁ H., BARTONIÈKA T., ORLOVA M. & VILÍMOVÁ J. 2012: Transport of bugs of the genus Cimex (Heteroptera: Cimicidae) by bats in western Palaearctic. Vespertilio 16: 43–54. BALVÍN O., VILÍMOVÁ J. & KRATOCHVÍL L. 2013: Batbugs (Cimex pipistrelli group, Heteroptera: Cimicidae) are morphologically, but not genetically differentiated among bat hosts. Journal of Zoological Systematics and Evolutionary Research 51: 287–295. BARTA M. 2008: Obrubnica západná, Leptoglossus occidentalis Heidemann, 1910 (Heteroptera: Coreidae), nový škodca ihliènatých drevín na Slovensku. (The western conifer seed bug, Leptoglossus occidentalis Heidemann, 1910 (Heteroptera: Coreidae), a new pest of in Slovakia). Pp. 307–314. In: Dendrologické dni v Arboréte Mlyòany SAV 2008. Zborník referátov z vedeckej konferencie. Days of dendrology in the Arboretum Mlynany SAS 2008. Proceedings of papers from scientific conference. Arborétum Mlyòany SAV, 15.–16. Október 2008, 379 pp (in Slovak, English summary). BARTA M. 2009: New facts about distribution and host spectrum of the invasive Nearctic conifer pest, Leptoglossus occidentalis (Heteroptera: Coreidae) in south-western Slovakia. Lesnícky Èasopis – Forestry Journal 55: 139–144. BARTONIÈKA T. 2010: Survival rate of bat bugs (Cimex pipistrelli, Heteroptera) under different microclimatic conditions. Parasitological Research 107: 827–833. BENEDEK P. 1970: The semiaquatic Heteroptera in the Carpathian basin with notes on the distribution and the phenology of the species. Faunistische Abhandlungen Staatliches Museum für Tierkunde in Dresden 3: 27–49. BEZUS’KO L. G. 1999: Poshirennya Ephedra distachya L. (Ephedraceae Wettst.) v pizn’ol’odovikiv’i ta golotseni na teritorii Ukraini (za palinologichnimi danimi). (Distribution of Ephedra distachya L. (Ephedraceae Wettst.) during the Late Glacial and Holocene on the territory of Ukraine (by palynological data). Ukrains’kyi Botanichnyi Zhurnal 56: 300–304 (in Ukrainian, Russian and English summaries). BODA P. 2006: A vízi- és vízfelszíni-poloskafauna váhltozásának hosszú távú vizsgálata a Bihari-sík vízfolyásaiban. (Realignment of the fauna of aquatic and semiaquatic Heteroptera: a long time research in water-courses of the Bihar-plain). Hydrológiai Közlöny 86: 18–20 (in Hungarian, English summary). BODA P., CSABAI Z., MÓRA A. & DÉVAI GY. 2004: Vízi- és vízfelszíni-poloskák (Heteroptera: Nepomorpha, Gerromorpha) mennyiségi elõfordulása egy alföldi mocsár két sásállományában. [Quantitative distribution of aquatic and semiaquatic bugs (Heteroptera: Gerromorpha, Nepomorpha) in two different sedge stands of a lowland marsh]. Hydrológiai Közlöny 84: 23–25 (in Hungarian, English summary without title). BODA P., VÁRBÍRÓ G. & DEÁK CS. 2012: Contribution to the aquatic macroinvertebrate fauna of some Hungarian water bodies. Acta Biologica Debrecina, Supplementum Oecologia Hungarica 28: 17–32. BOZDÌCHOVÁ J. 1973: Pøíspìvek k poznání fauny ploštic (Heteroptera) západních Èech. (Beitrag zur Kenntnis der Wanzenfauna (Heteroptera) von Westböhmen). Zprávy Muzejí Západoèeského Kraje, Pøíroda 15: 25–29 (in Czech, German summary). BRYJA J. & KMENT P. 2001: The present state of knowledge of the true bugs (Heteroptera) in the PLA of Poodøí (Czech Republic). Klapalekiana 37: 1–36. BRYJA J. & KMENT P. 2006a: Ploštice (Heteroptera) Chránìné krajinné oblasti Kokoøínsko. (True bugs (Heteroptera) of Kokoøínsko Protected Landscape Area). Bohemia Centralis 27: 267–294 (in Czech, English summary). BRYJA J. & KMENT P. 2006b: Ploštice (Heteroptera) pøírodní rezervace Svatý kopeèek a blízkého okolí (CHKO a BR Pálava, Èeská republika). (True bugs (Heteroptera) of the Svatý kopeèek Nature Reserve and its environs) (Pálava Protected Landscape Area and Biosphere Reserve, Czech Republic). Sborník Pøírodovìdeckého Klubu v Uherském Hradišti 8 (2003): 54–82 (in Czech, English summary). CARAPEZZA A. 2002: Heteroptera of Jordan: new taxa and new records. Naturalista Siciliano 26: 35–76. CASSIS G. & GROSS G. F. 1995: Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha). In: HOUSTON W. W. K. & MYANARD G. W. (eds.): Zoological Catalogue of Australia. Vol. 27.3A. CSIRO, Melbourne, xv + 506 pp. CASSIS G. & GROSS G. F. 2002: Hemiptera: Heteroptera (Pentatomomorpha). In: HOUSTON W. W. K. & WELLS A. (eds.): Zoological Catalogue of Australia. Vol. 27.3B. CSIRO, Melbourne, xiv + 737 pp.

528 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

COULIANOS C. C. 1989: Nya landskapsfynd an barkstinkflyn (Hem.-Het., Aradidae) jämte Aradus truncatus, ny för Sverige. (New provincial records of Swedish flatbugs and barkbugs (Hem.-Het., Aradidae) with Aradus truncatus Fieber, 1861, new to Sweden). Entomologisk Tidskrift 110: 53–57 (in Swedish, English summary). COULIANOS C. C. 2005: Annotated checklist and distribution of true bugs (Hemiptera-Heteroptera) of Estonia. Proceedings of the Estonian Academy of Sciences, Biology and Ecology 54: 136–165. CUPPEN J. G. M. 1988: Sigara iactans new for the Netherlands (Heteroptera: Corixidae). Entomologische Berichten (Amsterdam) 48: 94–96. DALLA TORRE C. W. VON 1877: Entomologische Notizen aus dem Egerlande. Jahres-Bericht des Natur- Historischen Vereins „Lotos“ 27: 91–208. DAVIDEANU A. 1999: Contribuþii la studiul heteropterelor acvatice din România. [Contribution to the study of aquatic Heteroptera of Romania]. Ph.D. Thesis, “Alexandru Ioan Cuza” University, Iaºi, 250 pp (in Romanian). DERJANSCHI V. & P ÉRICART J. 2006: Hémiptères Pentatomoidea Euro-Méditerranéens. Volume I. Faune de France. Vol. 90. Fédération Française des Sociétés de Sciences Naturelles, Paris, 494 pp + 16 pls. DOBŠÍK B. 1979: Synusie ploštic (Heteroptera Geocorisae Leston, Pendergrast, Southwood, 1954) bylinného patra Chránìného nalezištì Sedlisko. (Synusien der Wanzen (Heteroptera Geocorisae Leston, Pendergrast, Southwood, 1954) der Pflanzenschicht der geschützten Fundstätte Sedlisko). Západné Slovensko 6: 119–123 (in Czech, Russian and German summaries). DOBŠÍK B. 1988: Suchozemské ploštice (Heteroptera, Cimicomorpha, Pentatomomorpha) Plešivecké planiny. (Terrestrial bugs (Heteroptera, Cimicomorpha, Pentatomomorpha) of the Plešivecká planina Plateau). Ochrana Prírody 6B: 255–269 (in Czech; English, German and Russian summaries). DOBŠÍK B. 1991: Suchozemské ploštice (Heteroptera, Geocorisae) Zoborských vrchù. (Terrestrial bugs (Heteroptera, Geocorisae) of the Zobor Hills). Zobor 2: 55–66 (in Czech; Russian, German and English summaries). DOLLING W. R. 1972: A new species of Dicyphus Fieber (Hem., Miridae) from Southern England. Entomologist’s Monthly Magazine 107: 244–245. DOLLING W. R. 2006: Family Coreidae Leach, 1815. Pp. 43–101. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 5, Pentatomomorpha II. The Netherlands Entomological Society, Amsterdam, xiii + 550 pp. DUDA L. 1884: Soustavný pøehled èeského hmyzu polokøídlého (Hemiptera-Heteroptera). [Systematic synopsis of the Czech hemipterous insects (Hemiptera-Heteroptera)]. Pp. 1–39. In: KLUMPAR J. (ed.): Výroèní zpráva Cís. Král. Vyššího Gymnasia v Hradci Králové. Cís. Král. Vyšší Gymnasium v Hradci Králové, Hradec Králové (in Czech). DUDA L. 1885: Beiträge zur Kenntnis der Hemipteren-Fauna Böhmens. (7. Fortsetzung.). Wiener Entomologische Zeitung 4: 257–260. DURSUN A. & FENT M. 2013: Overview of the subgenus Ventocoris s. str. (Hemiptera: Heteroptera: Pentatomidae) with new records and a revised key to the Ventocoris species of Turkey. Zootaxa 3681: 151–177. DUTTO M. & CARAPEZZA A. 2011: Correlazione fra infestazioni domestiche da Arocatus melanocephalus (Hemiptera: Lygaeidae) e condizioni meteorologiche. Analisi di un caso in Piemonte. Il Naturalista Valtellinese – Atti Museo Civico di Storia Naturale di Morbegno 22: 65–69. DUWAL R. K., JUNG S.-H. & LEE S.-H. 2013: Resurrection of the genus Parapsallus Wagner (Hemiptera: Heteroptera: Miridae: Phylinae: Phylini). Zootaxa 3647: 597–600. DVOØÁK L., SYCHRA J. & DVOØÁKOVÁ K. 2010: Poznámky k entomofaunì bývalé pískovny u Pamferovy Huti (SZ Šumava). (Notes to the entomofauna of former sand-pit near Pamferova Huť (southwestern Bohemian Forest. Erica 17: 103–121. EHRENDORFER F. & HAMANN U. 1965: Vorschläge zu einer floristischen Kartierung von Mitteleuropa. Berichte der Deutschen Botanischen Gesellschaft 78: 35–50. ELDER J.-F. & CHÉRAU L. 2003: Une nouvelle espèce d’Hydrocorise pour la faune de France: Sigara (Subsigara) iactans Jansson, 1983 (Heteroptera, Corixidae). Bulletin de la Société Entomologique de France 108: 405–407. ESSER J. 2010: Ein Fund von Aradus bimaculatus Reuter, 1873 (Heteroptera, Aradidae) in Brandenburg nebst Bemerkungen zur Lebensweise der Art. Entomologische Nachrichten und Berichte 54: 141.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 529 P. K MENT ET AL.

FLOREN A. & GOGALA A. 2002: Heteroptera from beech (Fagus sylvatica) and silver fir (Abies alba) trees of the primary forest reserve Rajhenavski Rog, Slovenia. Acta Entomologica Slovenica 10: 25–32. GAO C.-Q., KONDOROSY E. & BU W.-J. 2013: A review of the genus Arocatus in the Palaearctic and Oriental Regions (Hemiptera: Heteroptera: Lygaeidae). Raffles Bulletin of Zoology 61: 687–704. GOGALA A. 2006: Heteroptera of Slovenia, III: Miridae. Annals for Istrian and Mediterranean Studies, Seria Historia Naturalis 16: 77–112. GOGALA A. & GOGALA M. 1986: Seznam vrst stenic, ugotovljenih v Sloveniji (Insecta: Heteroptera). (Check list of bug species recorded in Slovenia (Insecta: Heteroptera)). Biloški Vestnik 34: 21–52 (in Slovene, English abstract). GÖLLNER-SCHEIDING U. & ARNOLD K. 1988: Sammelausbeute von Heteropteren aus dem südwestlichen Bulgarien (Insecta). Faunistische Abhandlungen Staatliches Museum für Tierkunde Dresden 15: 137–154. GÜNTHER H., HOFFMANN H.-J., MELBER A., REMANE R., SIMON H. & WINKELMANN H. 1998: Rote Liste der Wanzen (Heteroptera). In: BUNDESAMT FÜR NATURSCHUTZ (eds.): Rote Listen gefährdeter Tiere Deutschlands. Schriftenreihe für Landschaftspglege und Naturschutz 55: 235–242. HARMUTH P. 2004: Kurze Mitteilung. P. 5. In: PFLANZENSCHUTZDIENST BADEN-WÜRTTEMBERG (ed.): Jahresbericht 2004. Stuttgart. HEISS E. 2001: Family Aradidae Brullé, 1836 – Flat Bugs. Pp. 3–34. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 4. Pentatomomorpha I. The Netherlands Entomological Society, Amsterdam, xiv + 346 pp. HEISS E. 2010: A new species and additional records of Aradus Fabricius, 1803 from China (Heteroptera, Aradidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 62: 47–54. HEISS E. & PÉRICART J. 2007: Hemiptères Aradidae, et Dipsocoromorphes Euro-Méditerranéens. Faune de France. Vol. 91. Fédération Française des Sociétés de Sciences Naturelles, Paris, 509 pp + 8 pls. HELIÖVAARA K. & VÄISÄINEN R. 1983: Enviromental changes and the flat bugs (Heteroptera, Aradidae and Aneuridae). Distribution and abundance in Eastern Fennoscandia. Annales Entomologici Fennici 49: 103–109. HENRY T. J. & WHEELER A. G. JR. 1976: Dicyphus rhododendri Dolling, first records from North America (Hemiptera: Miridae). Proceedings of the Entomological Society of Washington 78: 108–109. HENRY T. J. & WHEELER A. G. JR. 1988: Family Miridae Hahn, 1833 (= Capsidae Burmeister, 1835). The Plant Bugs. Pp. 251–507. In: HENRY T. J. & FROESCHNER R. C. (eds.): Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden, New York, Københaven, Köln, xix + 958 pp. HENRIKSON L. & OSCARSON H. G.1981: Corixids (Hemiptera – Heteroptera), the new top predators in acidified lakes. Verhandlungen Internationale Vereinigung für Theoretische und Angewandte Limnologie 21: 1616–1620. HENRIKSON L. & OSCARSON H. G. 1985: Waterbugs (Corixidae, Hemiptera – Heteroptera) in acidified lakes, habitat selection and adaptations. Ecological Bulletin 37: 232–238. HERCZEK A. & LUKÁŠ J. 1998: Mirid fauna (Heteroptera, Miridae) of the State Nature Reserve Devínska Kobyla in Slovakia. Acta Biologica Silesiaca 32(49): 13–18. HJÄLTÉN J., ATLEGRIM O., SANDSTRÖM F., PETTERSSON R. & REXSTAD E. A. 2006: Occurrence of flat bugs (Heteroptera: Aradidae) in burned and unburned forests. Entomologica Fennica 17: 130–135. HOBERLANDT L. 1944a: Heteroptera Bohemiae III. Acta Entomologica Musei Nationalis Pragae 21–22: 276–283 (in Czech and Latin). HOBERLANDT L. 1944b: Heteroptera Slovakiae. Èasopis Èeské Spoleènosti Entomologické 41: 23–31 (in Czech). HOBERLANDT L. 1944c: Aradus (Quilnus) mirus Bergr. v Èechách. [Aradus (Quilnus) mirus Bergr. In Bohemia]. Vìda Pøírodní 23(3): 91–92 (in Czech). HOBERLANDT L. 1947: Pøíspìvek k poznání Heteropter Èeskoslovenska VIII. (Heteroptera Moravy 4.) (Contribution to the knowledge of Heteroptera of Czechoslovakia VIII. (Heteroptera of Moravia 4.)). Èasopis Èeskoslovenské Spoleènosti Entomologické 44: 147–152 (in Czech, English summary). HOBERLANDT L. 1956: Aradus kuthyi Horváth zjištìný ve státní reservaci Kovaèovské kopce, nový pro faunu ÈSR. (Aradus kuthyi Horváth in the State Reserve Kovaèovské kopce – new for the fauna of Czechoslovakia). Acta Entomologica Musei Nationalis Pragae 30 (1955): 15–30 (in Czech and English).

530 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

HOBERLANDT L. 1963: Distributional notes on some species of Heteroptera from Czechoslovakia with a contribution on the taxonomy of the genus Hyoidea (Heteroptera, Miridae). Acta Faunistica Entomologica Musei Nationalis Pragae 9: 253–283. HOBERLANDT L. 1977a: Heteroptera. Pp. 61–82. In: DLABOLA J. (ed.): Enumeratio Insektorum Bohemoslovakiae. Acta Faunistica Entomologica Musei Nationalis Pragae, Supplementum 4: 1–158. HOBERLANDT L. 1977b: Distributional data on Saldidae (Heteroptera) in Czechoslovakia with a taxonomic note on Salda sahlbergi Reuter and Salda henschi (Reuter). Acta Entomologica Musei Nationalis Pragae 39: 139–158. HOFFMANN H.-J. 2010: Zum Vorkommen von Tupiocoris rhododendri (Dolling, 1972) (Heteroptera, Miridae) in NRW. Heteropteron 33: 31–32. HOFFMANN H.-J. & MELBER A. 2003: Verzeichnis der Wanzen (Heteroptera) Deutschlands. Pp. 209–272. In: KLAUSNITZER B. (ed.): Entomofauna Germanica. Vol. 6. Entomologische Nachrichten und Berichte, Beiheft 8: 1–344. HOFFMANN H.-J. & TERME L 2012: Zum Erstnachweis und Massenvorkommen der Ulmenwanze Arocatus melanocephalus (Fabricius, 1798) (Heteroptera, Lygaeidae) in Dortmund / Nordrhein-Westfalen. Heteropteron 38: 27–30. HRADIL K. 2010: Faunistic records from the Czech Republic – 290. Heteroptera: Miridae. Dicyphus escalerae Lindberg, 1934. Klapalekiana 46: 131–132. HRADIL K. 2011: Faunistic records from the Czech Republic – 315. Heteroptera: Lygaeidae. Belonochilus numenius (Say, 1831). Klapalekiana 47: 261–262. HRADIL K., KMENT P., BRYJA J., ROHÁÈOVÁ M., BAÒAØ P. & ÏURÈOVÁ K. 2008: New and interesting records of true bugs (Heteroptera) from the Czech Republic and Slovakia IV. Klapalekiana 44: 165–206 (in English, Czech summary). HRADIL K., PSOTA V. & Š TASTNÁ P. 2013: Species diversity of true bugs on apples in terms of plant protection. Plant Protection Science 49: 73–83. JANSSON A. 1995: Family Corixidae Leach, 1815 – water boatmen. Pp. 26–56. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1. Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha and Leptopodomorpha. The Netherlands Entomological Society, Amsterdam, xxvi + 222 pp. JEZIORSKI P., KMENT P., DITRICH T., STRAKA M., SYCHRA J. & DVOØÁK L. 2013: Distribution of Gerris asper and G. lateralis (Hemiptera: Heteroptera: Gerridae) in the Czech Republic. Klapalekiana 48(3–4) (2012): 191–202. JOSIFOV M. 1958: Heterocordylus cytisi n. sp. – eine neue Miriden-Art aus Bulgarien. Fragmenta Balcanica Musei Macedonici Scientiarum Naturalium 2: 83–89. JOSIFOV M. 1963: Polukrili nasekomi (Heteroptera) ot okolnostite na Petrich (yugozapadna B’lgariya). (Heteropteren aus der Umgebung von Petriè (SW Bulgarien).) Izvestiya na Zoologicheskiya Institut s Muzey (Sofia) 13: 93–132 (in Bulgarian, Russian and German summaries). JOSIFOV M. 1978: Dendrobiontni i dendrofilni polutv’rdokrili nasekomi (Heteroptera) po d’ba v B’lgariya. (Dendrobionte und dendrophile Halbflügler (Heteroptera) an der Eiche in Bulgarien.) Acta Zoologica Bulgarica 9: 3–14 (in Bulgarian, Russian and German summaries). JOSIFOV M. 1986: Verzeichnis der von der Balkanhalbinsel bekannten Heteropterenarten (Insecta, Heteroptera). Faunistische Abhandlungen, Staatliches Museum für Tierkunde Dresden 14: 61–93. JUHÁSZ P., KISS B., OLÁJOS P. & GRIGORSZKY I. 1999: Faunisztikai kutatások a Körös-Maros Nemzeti Park mûködési területén levõ “szentély” jellegû holtmedrekben. (Faunistical research on the ‘sanctuary’ oxbows of River Körös). Crisicum 2: 99–110 (in Hungarian, English summary). KÁLMÁN A., BODA R., KÁLMÁN Z., MAUCHART P., ROZNER GY., SZIVÁK I., SOÓS N. & CSABAI Z. 2011: Contribution to the aquatic macroinvertebrate fauna of the Zselic Hilly Region, SW Hungary. Acta Biologica Debrecina, Supplementum Oecologia Hungarica 26: 99–115. KANYUKOVA E. V. 2003: A key to species of Arctocorisa from Russia and Mongolia with description of A. germari mongolica ssp. n. (Heteroptera: Corixidae). Zoosystematica Rossica 11: 327–329. KANYUKOVA E. V. 2006: Vodnye poluzhestkokrylye nasekomye (Heteroptera: Nepomorpha, Gerromorpha) fauny Rossii i sopredelnykh stran. [Aquatic and semiaquatic (Heteroptera: Nepomorpha, Gerromorpha) of the fauna of Russia and neighbouring countries]. Dal’nauka, Vladivostok, 296 pp (in Russian, English abstract).

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 531 P. K MENT ET AL.

KANYUKOVA E. V. & VINOKUROV N. N. 2008: New data on distribution of bark bugs in Siberia and the Far East of Russia (Heteroptera: Aradidae). Zoosystematica Rossica 16(1) (2007): 48. KAPLIN V. G. 1993: Otkrytozhivushchie chlenistonogie sosudistykh rasteniy Garagum. [Openly living arthropods of vascular plants in Garagums]. Ylym, Ashgabat, 444 pp. (in Russian). KERZHNER I. M. 1964: Family Isometopidae. Family Miridae (Capsidae). Pp. 700–765. In: BEI-BIENKO G. YA. (ed.): Opredelitel’ nasekomykh evropeyskoy chasti SSSR v pyati tomakh. Vol. I. Nizhshie, drevnekrylye, s nepolnym prevrashcheniem. [Keys to the insects of the European part of the USSR in five volumes. Vol. 1. Apterygota, Palaeoptera, Hemimetabola.] Nauka, Moskva and Leningrad, 936 pp (in Russian). [English translation: 1967, Israel Program for Scientific Translation, Jerusalem, pp. 913–1003.] KERZHNER I. M. 1995: Infraorder Dipsocoromorpha. Pp. 6–12. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1, Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha and Leptopodomorpha. The Netherlands Entomological Society, Amsterdam, xxvi + 222 pp. KERZHNER I. M. & JOSIFOV M. 1999: Miridae Hahn 1833. Pp. 1–577. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 3. Cimicomorpha II. The Netherlands Entomological Society, Amsterdam, xiv + 577 pp. KLEMENTOVÁ B. 2012: Vodné bzdochy (Heteroptera) Slovenska: rozšírenie a ekológia. [Water bugs (Heteroptera) of Slovakia: distribution and ekology]. Diploma thesis. Fakulta ekológie a environmentalistiky, Technická univerzita vo Zvolene, Zvolen, 63 pp (unpublished). KLEMENTOVÁ B., MATÚŠOVÁ Z., NOVIKMEC M., OÈADLÍK M. & SVITOK M. 2011: Vodné bzdochy umelých vodných kanálov. [Aquatic true bugs of artificial water channels]. P. 16. In: STLOUKAL E. (ed.): Zborník abstraktov z konferencie 17. Feriancove dni 2011. Prírodovedecká fakulta UK, Bratislava, 24.–25.11.2011. Faunima, Bratislava, 34 pp. KLEMENTOVÁ B., SVITOK M. in press: Anisops sardeus (Heteroptera): a new expansive species in the Central Europe. Biologia (Bratislava). KLEMENTOVÁ B., SVITOK M., BITUŠÍK P., BULÁNKOVÁ E., KMENT P., MANKO P., MATÚŠOVÁ Z., NOVIKMEC M., OÈADLÍK M. & ROVNÝ F. 2012a: Vodné bzdochy Slovenska: rozšírenie a ekológia. (Water bugs of Slovakia: distribution and ecology). P. 63. In: ZAŤOVIÈOVÁ Z. (ed.): XVI. konferencia Slovenskej limnologickej spoloènosti a Èeské limnologické spoleènosti – Zborník príspevkov, 25.–29. jún 2012, Jasná. Slovenská limnologická spoloènosť pri SAV, Bratislava, 235 pp (in Slovak, English title). KLEMENTOVÁ B., SVITOK M. & GREGOR M. 2012b: Životný cyklus a produkcia vodných bzdôch Cymatia coleoptrata Fabricius 1777 a Plea minutissima Leach 1817. (Life cycle and secondary production of water bugs Cymatia coleoptrata Fabricius 1777 a Plea minutissima Leach 1817). P. 193. In: ÈIAMPOROVÁ- ZAŤOVIÈOVÁ Z. (ed.): XVI. konferencia Slovenskej limnologickej spoloènosti a Èeské limnologické spoleènosti – Zborník príspevkov, 25.–29. jún 2012, Jasná. Slovenská limnologická spoloènosť pri SAV, Bratislava, 235 pp (in Slovak, English title). KMENT P. 1999: Heteroptera. Pp. 187–194. In: OPRAVILOVÁ V., VAÒHARA J. & SUKOP I. (eds.): Aquatic invertebrates of the Pálava Biosphere Reserve of UNESCO. Folia Facultatis Scientiarum Naturalium Univiversitatis Masarykianae Brunensis, Biologica 101: 1–279. KMENT P. 2004: First records of two species of the genus Tuponia Reuter, 1875 (Heteroptera: Miridae) in Slovakia. Biológia (Bratislava) 59: 164. KMENT P. 2006: Heteroptera – ploštice. Pp. 255–266. In: MLÍKOVSKÝ J. & STÝBLO P. (eds.): Nepùvodní druhy fauny a flóry Èeské republiky. [Alien species of fauna and flora of the Czech Republic]. Èeský svaz ochráncù pøírody, Praha, 496 pp (in Czech). KMENT P. 2009: Èechy a Morava pro potøeby faunistického výzkumu. (Bohemia and Moravia for the purposes of faunistic research). Klapalekiana 45: 287–291 (in Czech, English summary). KMENT P. & BAÒAØ P. 2010: On the taxonomy and distribution of the genus Maccevethus (Hemiptera: Heteroptera: Rhopalidae). Acta Musei Moraviae, Scientiae Biologicae 95(1): 15–47. KMENT P. & BAÒAØ P. 2012: True bugs (Hemiptera: Heteroptera) of the Bílé Karpaty Protected Landscape Area and Biosphere Reserve (Czech Republic). Pp. 323–628. Acta Musei Moraviae, Scientiae Biologicae 96(2) (2011): 1–933. KMENT P. & BRYJA J. 2001: New and interesting findings of true bugs (Heteroptera) from the Czech Republic and Slovakia. Klapalekiana 37: 231–248.

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KMENT P., BRYJA J., JINDRA Z., HRADIL K. & BAÒAØ P. 2003: New and interesting records of true bugs (Heteroptera) from the Czech Republic and Slovakia II. Klapalekiana 39: 257–306. KMENT P., BRYJA J., HRADIL K. & JINDRA Z. 2005: New and interesting records of true bugs (Heteroptera) from the Czech Republic and Slovakia III. Klapalekiana 41: 157–213. KMENT P. & CUNEV J. in press: První nález nepùvodní ploštièky Belonochilus numenius (Hemiptera: Heteroptera: Lygaeidae) na Slovensku. (First record of the alien seed bug Belonochilus numenius (Hemiptera: Heteroptera: Lygaeidae) in Slovakia). Entomofauna Carpathica (submitted 2011) KMENT P. & DOLEJŠOVÁ K. 2010: The assassin bug Nagusta goedelii (Hemiptera: Heteroptera: Reduviidae) in Prague: an accidental introduction? Klapalekiana 46: 191–201. KMENT P., DVOØÁK M., HOVORKA O., KINDL J. & KRIST M. 2010: Faunistics records from the Czech Republic – 291. Heteroptera: Oxycarenidae. Oxycarenus lavaterae. Klapalekiana 46: 133–135. KMENT P., HRADIL K. & JINDRA Z. 2009b: The distribution of Pinthaeus sanguinipes in the Czech Republic and Slovakia (Hemiptera: Heteroptera: Pentatomidae). Acta Musei Moraviae, Scientiae Biologicae 94: 43–55. KMENT P. & J INDRA Z. 2008: New records of Eurydema fieberi from the Czech Republic with corrections to some previously published records of Palaearctic Eurydema species (Hemiptera: Heteroptera: Pentatomidae). Acta Musei Moraviae, Scientiae Biologicae 93: 11–27. KMENT P. & KEJVAL Z. 2011: První pøíspìvek k faunì ploštic (Hemiptera: Heteroptera) Èeského lesa. (First contribution to the true bug fauna (Hemiptera: Heteroptera) of the Èeský les Mts.). Klapalekiana 47: 29–53 (in Czech, English summary). KMENT P. & S MÉKAL A. 2002: Pøíspìvek k faunistice nìkterých vzácných vodních ploštic (Heteroptera: Nepomorpha, Gerromorpha) v Èeské republice. (Contribution to the faunistics of some rare water bugs (Heteroptera: Nepomorpha, Gerromorpha) in the Czech Republic). Sborník Pøírodovìdného Klubu v Uherském Hradišti 7: 155–181 (in Czech, English summary). KMENT P., ŠTYS P., EXNEROVÁ A., TOMŠÍK P., BAÒAØ P. & HRADIL K. 2009a: The distribution of Tropidothorax leucopterus in the Czech Republic and Slovakia (Hemiptera: Heteroptera: Lygaeidae). Acta Musei Moraviae, Scientiae Biologicae 94: 27–42. KMENT P. & V ILÍMOVÁ J. 2006: Heteroptera (ploštice). Pp. 139–146. In: FARKAÈ J., KRÁL D. & ŠKORPÍK M. (eds.): Èervený seznam ohrožených druhù Èeské republiky. Bezobratlí. Red list of threatened species in the Czech Republic. Invertebrates. Agentura ochrany pøírody a krajiny ÈR, Praha (2005), 760 pp (in Czech and English). KMENT P. & V ILÍMOVÁ J. 2010: Thoracic scent efferent system of Pentatomoidea (Hemiptera: Heteroptera): a review of terminology. Zootaxa 2706: 1–77. KNYSHOV A. & KONSTANTINOV F. V. 2013: A taxonomic revision of the genus Hyoidea (Hemiptera: Heteroptera: Miridae). Acta Entomologica Musei Nationalis Pragae 53: 1–32. KOLAØÍKOVÁ K., HORECKÝ J., STUCHLÍK E., LIŠKA M., JÍCHOVÁ M., TÁTOSOVÁ J., LAPŠANSKÁ N., HOØICKÁ Z., CHVOJKA P., BERAN L., KOŠEL V., MATÌNA J., ÈIAMPOROVÁ-ZAŤOVIÈOVÁ Z., KRNO I., BULÁNKOVÁ E., ŠPORKA F. & KMENT P. in press: Benthic macroinvertebrates along the Czech part of the Labe and lower section of the Vltava from 1996–2005 with a particular focus on rare and alien species. Biologia (Bratislava). KOLLÁR J., HRUBÍK P. & T KÁÈOVÁ S. 2009: Monitoring of harmful insect species in urban conditions in selected model areas of Slovakia. Plant Protection Science 45: 119–124. KONDOROSY E. 2000: Adatok a Villányi-hegység poloskanépességének (Heteroptera) ismeretéhez. (Data to the knowledge of the Heteroptera fauna of the Villány Hills (Hungary)). Dunántúli Dolgozatok Természettudományi Sorozat 10: 165–174 (in Hungarian, English summary). KONDOROSY E. 2001: Somogy megye poloskafaunája (Heteroptera). (Bug fauna of Somogy county (Heteroptera)). Natura Somogyensis 1: 123–134 (in Hungarian, English summary). KONDOROSY E. 2005: New true bug species in the Hungarian fauna (Heteroptera). Folia Entomologica Hungarica 66: 17–22. KONDOROSY E. 2011: Keszthely és környéke poloskafaunájának alapvetése (Heteroptera). (Heteroptera fauna of Keszthely and vicinity). Folia Musei Historico-Naturalis Bakonyiensis 28: 105–145 (in Hungarian, English abstract). KONDOROSY E. & FÖLDESSY M. 1998: Adatok a Duna-Dráva Nemzeti Park Dráva menti területei poloska (Heteroptera) faunájához. (Data to the Heteroptera fauna of areas along river Dráva in the Duna (Danube)- Dráva National Park). Dunántúli Dolgozatok Természettudományi Sorozat 9: 159–176 (in Hungarian, English summary).

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KONDOROSY E., SZÉL GY. & MERKL O. 1996: Adatok a Kis-Balaton poloska- és bogárfaunájához. (Data to the Heteroptera and Coleoptera fauna of the Kis-Balaton). Pp. 309–322. In: POMOGYI P. (ed.): 2. Kis-Balaton Ankét: Keszthely, 1996, szeptember 9–11.: összefoglaló értékelés a Kis-Balaton Védõrendszer 1991–1995 közötti kutatási eredményeirõl. Keszthely, 713 pp (in Hungarian). KRAJEWSKI S. 1969: Pluskwiaki wodne (Heteroptera) rzeki Grabi i jej terenu zalewowego. (Wasserwanzen (Heteroptera) des Flusses Grabia und seines Überschwemmungsgebieten). Polskie Pismo Entomologiczne 39: 465–513 (in Polish, German summary). KRIST M. & KMENT P. 2010: Nález kriticky ohroženého druhu ploštice Macrosaldula scotica (Curtis, 1835) (Heteroptera: Saldidae) na štìrkopískových lavicích v CHKO Litovelské Pomoraví s pøehledem o rozšíøení druhu v ÈR. (Discovery of critically endangered shore bug Macrosaldula scotica (Curtis, 1835) (Heteroptera: Saldidae) on gravel banks in Litovelské Pomoraví PLA with review of its distribution in the Czech Republic). Zprávy Vlastivìdného Muzea v Olomouci 299: 37–43 (in Czech, English abstract). KRIŠTOFÍK J. & DANKO Š. 2012: Arthropod ectoparasites (Acarina, Heteroptera, Diptera, Siphonaptera) of bats in Slovakia. Vespertillio 16: 167–189. KURZ¥TKOWSKA A. 1999: Water bugs (Hetemptera) of high bogs and transitional moors of Masurian Lake District. Polskie Pismo Entomologiczne 68: 349–369. LACINA A. 2010: Nález klopušky jesenické po 63 letech nezvìstnosti. (The discovery of Pithanus hrabei after disappearance of 63 years). Živa 58/96: 270 + civ (in Czech, separate English summary). LACINA A. 2011: Faunistic records from the Czech Republic – 311. Heteroptera: Miridae. Pithanus hrabei Stehlík, 1952. Klapalekiana 47: 101–102. LINDSKOG P. 1991: Taxonomy and distribution of the boreomontane shore bugs Salda sahlbergi and S. henschii (Heteroptera, Saldidae). Entomologisk Tidskrift 112: 1–18. LINDSKOG P. 1995: Infraorder Leptopodomorpha. Pp. 115–141. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1, Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha and Leptopodomorpha. The Netherlands Entomological Society, Amsterdam, xxvi + 222 pp. LINNAVUORI R. E., SARAFRAZI M. A. & HOSYNI R. 1998: On the Isometopus species of Iran (Heteroptera: Miridae, Isometopinae). Applied Entomology and Phytopatology 66: 6–10. LIS B. 2012: True-bugs (Hemiptera: Heteroptera) of the botanical garden in Štramberk in Moravia (Czech Republic). Heteroptera Poloniae, Acta Faunistica 4: 27–37. LIU H.-Y., LUO Z.-H., CAO L.-M. & CAI W.-ZH. 2011: Peirates hybridus (Scopoli), a new record species of the genus Peirates (Hemiptera: Reduviidae: Peiratinae) from China. Entomotaxonomia 33: 109–111. MAISTRELLO L., LOMBROSO L., PEDRONI E., REGGIANIA A. & VANIN S. 2006: Summer raids of Arocatus melanocephalus (Heteroptera, Lygaeidae) in urban buildings in Northern Italy: Is climate change to blame? Journal of Thermal Biology 31: 594–598. MALENOVSKÝ I., BAÒAØ P. & KMENT P. 2011: A contribution to the faunistics of Hemiptera (Cicadomorpha, Fulgoromorpha, Heteroptera, and Psylloidea) associated with dry grassland sites in southern Moravia (Czech Republic). Acta Musei Moraviae, Scientiae Biologicae 96(1): 41–187. MALENOVSKÝ I., KMENT P. & S YCHRA J. in press: Ploštice, køísi a mery (Hemiptera: Heteroptera, Auchenorrhyncha et Psylloidea) okolí Pøebuzi v Krušných horách. (True bugs, leafhoppers, planthoppers and psyllids (Hemiptera: Heteroptera, Auchenorrhyncha et Psylloidea) of the environs of Pøebuz in the Krušné hory Mountains (Czech Republic)). Klapalekiana. MANKO P. 2011: Interesuj¹ce stwierdzenia trzech rzadkich i zagro¿onych merolimnicznych gatunków owadów na S³owacji. (Interesting findings of three rare and endangered merolimnic insect species in Slovakia). Forum Faunistyczne 1(1): 56–62 (in Polish, English summary). MATOCQ A. 1991: Contribution à l’étude de genre Conostethus Fieber, avec la description d’une espèce nouvelle de France (Heteroptera, Miridae). Nouvelle Revue d’Entomologie (N. S.) 8: 135–148. MATOCQ A. & PLUOT-SIGWALT D. 2012: Revision des Amblytylus et essai de mise au point sur les genres Amblytylus Fieber et Megalocoleus Reuter (Heteroptera: Miridae: Phylinae). Annales de la Société Entomologique de France (Nouvelle Serie) 48: 123–154. MELBER A., GÜNTHER H. & RIEGER CH. 1991: Die Wanzenfauna des Österreichischen Neusiedlerseegebietes (Insecta, Heteroptera). Wissenschaftlichen Arbeiten aus dem Burgenland 89: 63–192. MÓRA A., DEÁK CS., KÁLMÁN A., KÁLMÁN Z., LÕKKÖS A., SOÓS N. & CSABAI Z. 2011: Contribution to the aquatic insect fauna of Káli-Medence and Fekete-Hegy, and their surroundings (Balaton Uplands). Folia Musei Historico-Naturalis Bakonyiensis 28: 147–180.

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MORKEL C. 2006: Erste Nachweise der Blindwanze Phytocoris hirsutulus (Flor, 1861) (Insecta: Heteroptera, Miridae) in Hessen. Philippia 12: 361–363. MOULET P. 1995: Hémiptères Euro-Méditerranéens. Faune de France. Vol. 81. Fédération Française des Sociétés de Sciences Naturelles, Paris, iv + 336 pp + 3 pls in text. NAU B. S. & BROOKE S. E. 2006: Two water bugs new to Britain, Cymatia rogenhoferi Fieber and Sigara iactans Jansson (Hem., Corixidae), with comments on migration. Entomologist’s Monthly Magazine 142: 229–234. NICKERL O. SEN. 1905: Beiträge zur Insekten-Fauna Böhmens. II. Fundorte böhmischer Wanzenarten, nach der vom † MUDr. Ottokar Nickerl jun. hinterlassenen Hemipterensammlung zusammengestellt. Verlag der Gesellschaft für Physiokratie in Böhmen, Prag, iv + 43 pp. NOSEK J. N., VÁSÁRHELYI T., BAKONYI G. & OERTEL N. 2007: Spatial pattern of water bugs (Nepomorpha, Gerromorpha) at different scales in the Szigetköz (Hungary). Biologia (Bratislava) 62: 345–350. NOVÁK I. 1989: Seznam lokalit a jejich kódù pro síťové mapování entomofauny Èeskoslovenska. [List of localities and their grid codes used in faunistic mapping in Czechoslovakia]. Zprávy Èeskoslovenské Spoleènosti Entomologické pøi ÈSAV 25: 3–84 (in Czech). ORSZÁGH I. 1966: Príspevok k poznaniu fauny bzdôch (Heteroptera) Jurského šúru a pri¾ahlých svahoch Malých Karpát. (Contribution to further knowledge of Heteroptera fauna of „Jurský šùr“ and the adjacent slopes of the Little Carpathians). Entomologické Problémy 6: 37–75 (in Slovak, Russian, English and German summaries). ORTVAY T. 1902: XIV. Fejezet. Az izeltlábúak köre. A félszárnyúak. [Chapter XIV. Circle of the arthropods. The Hemiptera]. Pp. 510–558. In: ORTVAY T.: Pozsonyvármegye és területén fekvö Pozsony, Nagyszombat, Bazin, Modors, Szentgyörgy városok állatvilága. Elsö kötet: Állatrajzi rész. [Fauna of Bratislava County and cities of Bratislava, Trnava, Pezinok, Modrá, Svätý Jur of its territory. Vol. 1: Zoographic part]. Stampfel Károly, Pozsony [= Bratislava], 648 pp (in Hungarian). OSSIANNILSSON F. 1967: Anmärkningar och tillägg till Sveriges heteropterfauna (Hem.). (Notes on and additions to the Swedish fauna of Heteroptera (Hem.)). Entomologisk Tidskrift 88: 83–86 (in Swedish, English summary). PAPÁÈEK M. 1991: K bionomii znakoplavek (Heteroptera, Notonectidae) jižních a západních Èech. (On the bionomy of the backswimmers (Heteroptera, Notonectidae) from South and West Bohemia). Sborník Jihoèeského Muzea v Èeských Budìjovicích, Pøírodní Vìdy 31: 23–28 (in Czech, English summary). PAPÁÈEK M. 2002: K rozšíøení a charakteristice habitatù pìti druhù relativnì vzácných vodních ploštic (Heteroptera: Nepomorpha) v jihozápadním pøíhranièí Èeské republiky. (On the distribution and habitat preference of five relatively rare water bug species (Heteroptera: Nepomorpha) in the southwest border area of the Czech Republic). Pp. 225–231. In: PAPÁÈEK M. (ed.): Biodiverzita a pøírodní podmínky Novohradských hor. Sborník pøíspìvkù z konference 10. a 11. ledna 2002. Jihoèeská univerzita v Èeských Budìjovicích a Entomologický ústav AV ÈR, Èeské Budìjovice, 285 pp (in Czech, English summary). PAPÁÈEK M. 2004: Vodní ploštice (Hemiptera: Heteroptera: Nepomorpha, Gerromorpha). [Water bugs (Hemiptera: Heteroptera: Nepomorpha, Gerromorpha)]. Pp. 126–135. In: PAPÁÈEK M. (ed.): Biota Novohradských hor: modelové taxony, spoleèenstva a biotopy. (Biota of the Novohradské hory Mts.: Model taxa, communities and biotopes). Jihoèeská univerzita, Èeské Budìjovice, 304 pp (in Czech). PÉRICART J. 1965: Contribution a la faunistique de la Corse: Heteropteres Miridae et Anthocoridae. Bulletin Mensuel de la Société Linnéene de Lyon 34: 377–384. PÉRICART J. 1983: Hémiptères Tingidae Euro-Méditerranéens. Faune de France et régiones limitrophes. Vol. 69. Fédération Française des Sociétés de Sciences Naturelles, Paris, 618 pp. PÉRICART J. 1999a: Hémiptères Lygaeidae Euro-Méditerranéens. Vol. 1. Faune de France et régiones limitrophes. Vol. 84A. Fédération Française des Sociétés de Sciences Naturelles, Paris, xx + 468 pp + 6 pls. PÉRICART J. 1999b: Hémiptères Lygaeidae Euro-Méditerranéens. Vol. 2. Faune de France et régiones limitrophes. Vol. 84B. Fédération Française des Sociétés de Sciences Naturelles, Paris, iii + 453 pp + 3 pls. PÉRICART J. 2001: Family Lygaeidae Schilling, 1829 – Seed-bugs, pp. 35-220. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 4. Pentatomomorpha I. The Netherlands Entomological Society, Amsterdam, xiv + 346 pp. PÉRICART J. 2010: Hémiptères Pentatomoidea Euro-Méditerranéens. Vol. 3. Podopinae et Asopinae. Faune de France. Vol. 93. Fédération Française des Sociétés de Sciences Naturelles, Paris, 291 pp. + 24 pls. PÉRICART J. & GOLUB V. B. 1996: Tingidae Laporte, 1832 – lacebugs. Pp. 3–77. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 2. Cimicomorpha I. The Netherlands Entomological Society, Amsterdam, xiv + 361 pp.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 535 P. K MENT ET AL.

PERSSON A. 1966: Sigara fallenoidea (Hungerford) – en för Sverige ny Corixid – jämte några andra sällsyntare Heteroptera funna i Jokkmokkstrakten. [Sigara fallenoidea (Hungerford) – a new corixid for Sweden – as well as some other rare Heteroptera found in Jokkmokk region]. Entomologisk Tidskrift 87: 72–75 (in Swedish, English summary without title). POLHEMUS J. T. 1995: Family Notonectidae Latreille, 1802 – backswimmers. Pp. 63–73. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Vol. 1. Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha and Leptopodomorpha. The Netherlands Entomological Society, Amsterdam, xxvi + 222 pp. PROTIÆ LJ. 1998: Catalogue of the Heteroptera fauna of Yougoslav countries. Part one. Prirodnjaæki Muzej u Beogradu, Prosebna Izdanja 38: 1–215. PROTIÆ LJ. 2008: Contribution to the knowledge of the Isometopinae (Heteroptera, Miridae) of the Balkan Peninsula. Pp. 255–259. In: GROZEVA S. & SIMOV N. (eds.): Advances in Heteroptera Research. Festschrift in Honour of 80th Anniversary of Michail Josifov. Pensoft Publishers, Sofia-Moscow, 417 pp. PRUNER L. & MÍKA P. 1996: Seznam obcí a jejich èástí v Èeské republice s èísly mapových polí pro síťové mapování fauny. (List of settlements in the Czech Republic with associated map field codes for faunistic grid mapping system). Klapalekiana 32 (Supplementum): 1–115 (in Czech, English summary). PUTSHKOV P. V. 1987: Poluzhestkokrylye. Khishchnecy. [True bugs. Reduviidae]. Fauna Ukrainy 21(5). Naukova Dumka, Kiiv, 247 pp (in Russian). PUTSHKOV P. V. & M OULET P. 2010: Hémiptères Reduviidae d’Europe occidentale. Fédération Française des Sociétés de Sciences Naturelles, Paris, (2009), 668 pp + 24 pls. PUTSHKOV P. V. & P UTSHKOV V. G. 1996: Family Reduviidae Latreille, 1807 – assassin-bugs. Pp. 148–265. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearcic Region. Vol. 2. Cimicomorpha I. The Netherlands Entomological Society, Amsterdam, xiv + 361 pp. PUTSHKOV V. G. 1969: Ligeidi. [Lygaeidae]. Fauna Ukrainy, Vol. 21(3). Vidavnitstvo Akademii Nauk Ukrains’koi RSR, Kiiv, 388 pp (in Ukrainian). PUTSHKOV V. G. & PUTSHKOV P. V. 1983: Maloizvestnye poluzhestkokrylye (Heteroptera) Yuga SSSR. [Little known true bugs (Heteroptera) of the south of the USSR]. Vestnik Zoologii 1983(3): 17–25 (in Russian, English summary without title). RABITSCH W. 1999a: Die Wanzensammlung (Insecta, Heteroptera) von Johann Moosbrugger (1878–1953) am Naturhistorischen Museum Wien. Annalen des Naturhistorischen Museums Wien 101B: 163–199. RABITSCH W. 1999b: Neue und seltene Wanzen (Insecta, Heteroptera) aus Wien und Niederösterreich. Linzer Biologische Beiträge 31: 993–1008. RABITSCH W. 2002: Die Wanzenfauna (Heteroptera) der Sandberge bei Oberweiden im Marchfeld (Niederösterreich). Beiträge zur Entomofaunistik 3: 141–174. RABITSCH W. 2003: Neue und seltene Wanzen (Insecta, Heteroptera) aus Niederösterreich und Wien. Teil 3. Linzer Biologische Beiträge 35: 1293–1305. RABITSCH W. 2007: Rote Listen ausgewählter Tiergruppen Niederösterreichs – Wanzen (Heteroptera). 1 Fassung 2005. Amt der NÖ Landesregierung, Abteilung Naturschutz & Abteilung Kultur und Wissenschaft, St. Pölten, 280 pp. RABITSCH W. 2008: The times they are a-changin’ driving forces of recent additions to the Heteroptera fauna of Austria. Pp. 309–326. In: GROZEVA S. & SIMOV N. (eds.): Advances in Heteroptera research. Festschrift in honour of 80th anniversary of Michail Josifov. PenSoft Publishers, Sofia, Moscow, 417 pp. RABITSCH W. 2010a: Heteroptera. In: ROQUES A., KENIS M., LEES D., LOPEZ-VAAMONDE C., RABITSCH W., RASPLUS J.-Y. & ROY D. (eds.): Alien terrestrial arthropods of Europe. BioRisk 4: 407–433. RABITSCH W. 2010b: Harpocera hellenica Reuter, 1876 (Heteroptera: Miridae) – Neu für Österreich. Beiträge zur Entomofaunistik 11: 93–96. RAMPAZZI F. & DETHIER M. 1997: Gli eterotteri (Insecta: Heteroptera) delle torbiere a sfagni del Cantone Ticino e del Moesano (Val Calanca e Val Mesolcina – GR), Svizzeria. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 70: 419–439 (in Italian, English summary). REGGIANI A., PEDRONI E. & MAISTRELLO L. 2005: Infestazioni da Arocatus melanocephalus (Fabricius, 1798) (Heteroptera, Lygeidae) nella città di Modena. Atti della Società dei Naturalisti e Matematici di Modena 136: 119–125. RIDER D. A. 2006: Family Pentatomidae Leach, 1815. Pp. 233–402. In: AUKEMA B. & RIEGER CH. (eds.): Catalogue of the Heteroptera of the Palaearctic Region. Volume 5. Pentatomomorpha II. The Netherlands Entomological Society, Amsterdam, xiii + 550 pp.

536 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

RIETSCHEL S. 2000: Funde von Isometopus mirificus Mulsant & Rey, 1878 und Cardiastethus fasciiventris (Garbiglietti, 1869) in Oberrhein-Gebiet (Heteroptera, Isometopidae und Anthocoridae). Heteropteron 9: 9–10. ROHÁÈOVÁ M. 2008a: Structure and seasonal dynamics of the true bug assemblages (Heteroptera) on alien goldenrods Solidago spp. Èasopis Slezského Zemského Muzea (A) 57: 97–116. ROHÁÈOVÁ M. 2008b: Sludge-drying bed as a refuge of the endangered species. Scripta Facultatis Rerum Naturalium Universitatis Ostraviensis 186: 364–367. ROHÁÈOVÁ M. 2011: Ploštice (Heteroptera) beskydských a podbeskydských rašelinných biotopù 1. Pøírodní památka Obidová. (True bugs (Heteroptera) of the Beskidian and Subbeskidian peaty habitats 1. Obidová Nature Monument). Acta Musei Beskidensis 3: 85–102 (in Czech, English summary). ROHÁÈOVÁ M. 2012a: Ploštice (Heteroptera) pasínku Losový v Huslenkách (Vsetínské vrchy). (True bugs (Heteroptera) of traditionally grazed pasture „Losový“ in Huslenky (Vsetínské vrchy hills)). Acta Carpathica Occidentalis 2 (2011): 72–77 (in Czech, English summary). ROHÁÈOVÁ M. 2012b: Ploštice (Heteroptera) beskydských a podbeskydských rašelinných biotopù 2. Pøírodní rezervace Vøesová stráò. (True bugs (Heteroptera) of the Beskidian and Subbeskidian peaty habitats 2. Vøesová stráò Nature Reserve). Acta Musei Beskidensis 4: 155–172 (in Czech, English summary). ROHÁÈOVÁ M. 2013: Ploštice (Heteroptera) beskydských a podbeskydských rašelinných biotopù 3. Pøírodní rezervace Rybníky. (True Bugs (Heteroptera) of the Beskidian and Subbeskidian peaty habitats 3. Rybníky Nature Reserve). Acta Musei Beskidensis 5: in press (in Czech, English summary). ROHÁÈOVÁ M. & DROZD P. 2009: How many heteropteran species can live on alien goldenrods Solidago canadensis and S. gigantea in Europe? Biologia (Bratislava) 64: 981–993. ROSHKO G. M. 1976: Klopy-slepnyaki (Heteroptera, Miridae) Ukrainskikh Karpat, Zakarpat’ya i Prikarpat’ya. (Miridae (Heteroptera) of Ukrainian Karpathians [sic!]). Entomologicheskoe Obozrenie 55: 814–819 (in Russian, English abstract). ROUBAL J. 1957a: Studie o plošticích ze severozápadních Èech s kritickými poznámkami. (Eine Studie über Heteropteren von Nordwestböhmen nebst kritischen Bemerkungen). Acta Societatis Entomologicae Èechosloveniae 53: 63–109 (in Czech, Russian and German summaries). ROUBAL J. 1957b: Monografie èeských klešťanek (Corixidae). (Monographie der Ruderwanzen (Corixidae) Böhmens). Rozpravy ÈSAV, Øada Matematicko-Pøírodovìdná 67(9): 1–66 (in Czech, Russian and German summaries). ROUBAL J. 1958: Aradus bimaculatus Reut. (Heteroptera, Aradidae) v Èechách. (Aradus bimaculatus Reut. (Heteroptera, Aradidae) in Böhmen). Acta Entomologica Musei Nationalis Pragae 32: 279–284 (in Czech and German). ROUBAL J. 1959: O krkonošských Heteropterách. (De Heteropteris Asciburgii montis). Acta Musei Reginaehradecensis, Series A: Scientiae Naturales 2: 183–192 (in Czech, Latin summary). ROUBAL J. 1964: Heteroptera. P. 6. In: Entomofauna Tøebíèska. [Entomofauna of Tøebíè district]. Klub Pøírodovìdecký pøi Moravském Museu v Brnì, Brno (in Czech). ROUBAL J. 1968: K faunì Heteropter moravské hadcové stepi u Mohelna. (Zur Wanzenfauna des mährischen Serpentinsteppe bei Mohelno). Acta Musei Silesiae, Series A 18: 175–180 (in Czech, German summary). RUS I. 2005: Katalog sbírky ploštic (Heteroptera) kolínského rodáka Otokara Kubíka uložené v Regionálním muzeu v Kolínì – èást I. (Catalogue of the Heteroptera collection of Otokar Kubík deposited in Regional Museum Kolín. – Part I.). Práce Muzea v Kolínì, Øada Pøírodovìdná 6 (2004): 15–80 (in Czech, English summary). RUS I. & KMENT P. 2007: Katalog sbírky ploštic (Heteroptera) kolínského rodáka Otokara Kubíka uložené v Regionálním muzeu v Kolínì – èást II. (Catalogue of the Heteroptera collection of Otokar Kubík deposited in Regional Museum Kolín. – Part II.). Práce Muzea v Kolínì, Øada Pøírodovìdná 7 (2006): 47–80 (in Czech, English summary). SADÍLEK D., ŠŤÁHLAVSKÝ F., VILÍMOVÁ J. & ZIMA J. 2013: Extensive fragmentation of the X chromosome in the bed bug Cimex lectularius Linnaeus, 1758 (Heteroptera, Cimicidae): a survey across Europe. Comparative Cytogenetics 7: 253–269. SAVAGE A. A. 1989: Adults of the British aquatic Hemiptera Heteroptera. A key with ecological notes. Titus Wilson & Son, Kendal, 173 pp. SCHMITZ A., SCHÄTZEL H. & SCHMITZ H. 2010: Distribution and functional morphology of photomechanic infrared sensilla in flat bugs of the genus Aradus (Heteroptera, Aradidae). Arthropod Structure and Development 39: 17–25.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 537 P. K MENT ET AL.

SCHRAMEYER K. 2004: Foto mit Legende. Nachrichten aus der Deutschen Gesellschaft fü Allgemeine und Angewandte Entomologie 18(3): titel photo + 82. SEIDENSTÜCKER G. 1961: Heteropteren aus Bayern. Nachrichtenblatt der Bayerischen Entomologen 10: 12–16. SIMON H. 1992: Vergleichende Untersuchungen zur Wanzenfauna (Heteroptera) von Streuobstwiesen in Nordpfälzer Bergland. Beiträge zur Landespflege in Rheinland-Pfalz 15: 189–276. SIMON H. 2002: Erstes vorläufiges Verzeichnis der Wanzen (Insecta: Heteroptera) in Rheineland-Pfalz. Fauna Flora Rheinland-Pfalz 9(4): 1379–1420. SKALICKÁ A. 1995: Chamaecytisus Link – èilimník. Pp. 339–348. In: SLAVÍK B. (ed.): Kvìtena Èeské republiky. 4. (Flora of the Czech Republic. 4.) Academia, Praha, 529 pp (in Czech). SMITH C. L. 1988: Family Veliidae Amyot and Serville, 1843. The Small Water Striders. Pp. 734–742. In: HENRY T. J. & FROESCHNER R. C. (eds.): Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. E. J. Brill, Leiden, New York, Københaven, Köln, xix + 958 pp. SOLDÁN T., BOJKOVÁ J., VRBA J., BITUŠÍK P., CHVOJKA P., PAPÁÈEK M., PELTANOVÁ J., SYCHRA J. & TÁTOSOVÁ J. 2012: Aquatic insects of the Bohemian Forest glacial lakes: Diversity, long-term changes, and influence of acidification. Silva Gabreta 18: 123–283. SOÓS N., HORVAI V., CZIROK A. & CSABAI Z. 2009: Contribution to the aquatic and semiaquatic Heteroptera (Nepomorpha, Gerromorpha) fauna of the Transdanubian Region, Hungary. Acta Biologica Debrecensis, Supplementum Oecologica Hungarica 20: 193–208. SPITZER L., KONVIÈKA O., TROPEK R., ROHÁÈOVÁ M., TUF I. H. & NEDVÌD O. 2010: Spoleèenstvo èlenovcù (Arthropoda) zimujících na jedli bìlokoré (Abies alba) na Valašsku (okr. Vsetín, Èeská republika). (Assemblage of overwintering arthropods on white fir (Abies alba) in the Moravian Wallachia region (West Carpathians, Czech Republic). Èasopis Slezského Muzea Opava (A) 59: 217–232 (in Czech, English summary). STEHLÍK J. L. 1946: 12 nových druhù ploštic pro Moravu. [Twelve species of Heteroptera new for Moravia]. Entomologické Listy (Brno) 9: 143–145 (in Czech). STEHLÍK J. L. 1952: Fauna Heteropter Hrubého Jeseníku. (The fauna of Heteroptera of the mountain High Jeseník). Acta Musei Moraviae, Scientiae Naturales 37: 132–248 (in Czech, English and Russian summaries). STEHLÍK J. L. 1961: Pøíspìvek k poznání klopušek Moravy a Slovenska (Het. Miridae). (Beitrag zur Kenntnis der Blindwanzen Mährens und der Slowakei). Acta Musei Moraviae, Scientiae Naturales 46: 175–186 (in Czech, German summary). STEHLÍK J. L. 1985: Results of the investigations on Hemiptera in Moravia made by the Moravian Museum. (Pentatomoidea 4.). Acta Musei Moraviae, Scientiae Naturales 70: 147–175. STEHLÍK J. L. 1986: Results of the investigations on Hemiptera in Moravia made by the Moravian Museum. (Pentatomoidea 5). Acta Musei Moraviae, Scientiae Naturales 71: 147–178. STEHLÍK J. L. 1988: Results of the investigation on Hemiptera in Moravia made by the Moravian Museum (Coreoidea I.). Acta Musei Moraviae, Scientiae Naturales 73: 169–211. STEHLÍK J. L. & HEISS E. 2000: Results of investigations of the Hemiptera in Moravia made by the Moravian Museum (Aradidae, Pyrrhocoridae). Acta Musei Moraviae, Scientiae Biologicae 85: 333–350. STEHLÍK J. L. & HEISS E. 2001: Results of investigations on Heterptera in Slovakia made by the Moravian Museum (Aradidae, Pyrrhocoridae). Acta Musei Moraviae, Scientiae Biologicae 86: 177–194. STEHLÍK J. L. & HOBERLANDT L. 1953: Zajímavé nálezy Heteropter na Moravì a na Slovensku. (Interesting founds of Heteroptera in Moravia and Slovakia). Acta Musei Moraviae, Scientiae Naturales 38: 160–167 (in Czech, Russian and English summaries). STEHLÍK J. L. & ŠTEPANOVIÈOVÁ O. 1961: Pøedbìžná zpráva o výzkumu Heteropter vátých pískù na jižním Slovensku. (Vorbericht über die Erforschung von Heteropteren der Flugsande in der Südslowakei). Acta Musei Moraviae, Scientiae Naturales 46: 171–173 (in Czech, German summary). STEHLÍK J. L. & VAVØÍNOVÁ I. 1993: Results of the investigations on Heteroptera in Slovakia made by the Moravian Museum (Pentatomoidea II). Acta Musei Moraviae, Scientiae Naturales 77: 157–208. STEHLÍK J. L. & VAVØÍNOVÁ I. 1994: Results of the investigations on Heteroptera in Slovakia made by the Moravian Museum. (Pentatomoidea III). Acta Musei Moraviae, Scientiae Naturales 78 (1993): 99–163. STEHLÍK J. L. & VAVØÍNOVÁ I. 1995: Results of the investigations on Heteroptera in Slovakia made by the Moravian Museum. (Stenocephalidae, Coreidae, Alydidae, Rhopalidae). Acta Musei Moraviae, Scientiae Naturales 79 (1994): 97–147.

538 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

STEHLÍK J. L. & VAVØÍNOVÁ I. 1996: Results of the investigations on Heteroptera in Slovakia made by the Moravian Museum (Lygaeidae I). Acta Musei Moraviae, Scientiae Naturales 80 (1995): 163–233. STEHLÍK J. L. & VAVØÍNOVÁ I. 1997a: Results of the investigations on Hemiptera in Moravia made by the Moravian Museum (Reduviidae, Phymatidae, Nabidae: Prostemmatinae). Acta Musei Moraviae, Scientiae Naturales 81 (1996): 205–229. STEHLÍK J. L. & VAVØÍNOVÁ I. 1997b: Results of the investigations on Hemiptera in Moravia made by the Moravian Museum (Lygaeidae I). Acta Musei Moraviae, Scientiae Naturales 81 (1996): 231–298. STEHLÍK J. L. & VAVØÍNOVÁ I. 1998: Results of the investigations on Heteroptera in Slovakia made by the Moravian Museum (Reduviidae, Phymatidae, Nabidae: Prostemmatinae). Acta Musei Moraviae, Scientiae Biologicae 82 (1997): 109–126. STRAKA M., KMENT P., SYCHRA J. & HELEŠIC J. 2009: The proposed Úvalský rybník Nature Monument, an important refuge for wetland insects in South Moravia (Czech Republic): A species inventory of Odonata, Heteroptera and Coleoptera (partim) with the first Czech record of Corixa panzeri (Corixidae). Acta Musei Moraviae, Scientiae Biologicae 94: 87–116. SUCHÁ P. 2010: Seasonal dynamics in corixids (Heteroptera: Corixidae): A study from managed fishponds in South Bohemia. Acta Phytopathologica et Entomologica Hungarica 45: 107–114. SVENSSON B. G., TALLMARK B. & PETERSSON E. 2000: Habitat heterogenity, coexistence and habitat utilization in five backswimmer species (Notonecta spp.; Hemiptera, Notonectidae). Aquatic Insects 22: 81–98. SYCHRA J. & KMENT P. 2009: Vodní ploštice (Heteroptera: Nepomorpha, Gerromorpha) Rolavských vrchovišť. Aquatic bugs (Heteroptera: Nepomorpha, Gerromorpha) of Rolavská vrchovištì raised bogs. Pøiroda Kraslicka 2: 135–156. (in Czech, English abstract.) ŠTEPANOVIÈOVÁ O. 1977: Zmeny v spoloèenstvách bzdôch (Heteroptera) Blatskej nížiny vyvolané stavbou Zemplínskej šíravy. Acta Facultatis Rerum Naturalium Universitatis Comenianae, Zoologica 22: 45–68 (in Slovak, Russian summary). ŠTEPANOVIÈOVÁ O. 1991: Formation of taxocoenoses of Heteroptera in plant communities of relatively unaffected ecosystems and those affected by human activities. Acta Facultatis Rerum Naturalium Universitatis Comenianae, Zoologica 35: 31–51. ŠTEPANOVIÈOVÁ O. 2001: Bzdochy (Heteroptera) lesostepních biotopov Prírodnej rezervácie Kòaží [sic!] vrch v Tematínskych kopcoch (Povážský Inovec). (Bugs (Heteroptera) in the woody steppes habitats of the Nature Rezerve [sic!] Kòazí vrch in Tematínske kopce (Považský Inovec Mts)). Folia Faunistica Slovaca 6: 75–79 (in Slovak, English abstract). ŠTEPANOVIÈOVÁ O. 2003: First record of Arocatus melanocephalus (Heteroptera, Lygaeidae) from Slovakia. Entomological Problems 33: 30. ŠTEPANOVIÈOVÁ O. & BIANCHI Z. 2001: Èervený (ekosozologický) zoznam bzdôch (Heteroptera) Slovenska. (Red (ecosozological) list of Heteroptera of Slovakia). In: BALÁŽ D., MARHOLD K. & URBAN P. (eds.), Èervený zoznam rastlín a živoèíchov Slovenska. (Red list of plants and animals of Slovakia). Ochrana Prírody 20(Suppl.): 105–106 (in Slovak, English summary). ŠTEPANOVIÈOVÁ O. & BIANCHI Z. 2003: K faune bzdôch (Heteroptera) viatych pieskov Podunajskej nížiny. (On the fauna of bugs (Heteroptera) of blown sands in the Podunajská bížina lowland). Entomofauna Carpathica 15: 35–40 (in Slovak, English abstract). ŠTUSÁK J. M. 1976: Faunistics records from Czechoslovakia. Heteroptera. Lygaidae, Tingidae, Aradidae. Acta Entomologica Bohemoslovaca 73: 430. ŠTUSÁK J. M. 1980: Øád Ploštice – Heteroptera. [Order True bugs – Heteroptera]. Pp. 133–155. In: ROZKOŠNÝ R. (ed.): Klíè vodních larev hmyzu. [Key to aquatic larvae of insects]. Academia, Praha, 521 pp (in Czech). ŠTYS P. 1960: The Czechoslovakian populations of Notonecta reuteri Hungerford, 1928 (Het., Notonectidae). Èasopis Èeskoslovenské Spoleènosti Entomologické 57: 129–135. ŠTYS P. 1961: Die Wanzenfauna des Moorgebietes Soos in Böhmen (Heteroptera). Acta Universitatis Carolinae Biologica, Supplementum 1960: 83–133. ŠTYS P. 1972: Hojný výskyt chvojníku dvouklasého v Èenkovì mimo území státní pøírodní rezervace. [Abundant occurrence of Ephedra distachya in Èenkov outside the territory of the State Nature Reserve]. Ochrana Pøírody 27(2): 41 (in Czech). ŠTYS P. 1976: Faunistic records from Czechoslovakia. Heteroptera. Corixidae, Tingidae, Aradidae. Acta Entomologica Bohemoslovaca 73: 61.

Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 539 P. K MENT ET AL.

ŠTYS P. & Š KAPEC L. 1992: Klopuška bzdôška Hyoidea notaticeps Reuter, 1875. Pp. 74–75. In: ŠKAPEC L. (ed.): Èervená kniha ohrožených a vzácných druhù rostlin a živoèichù ÈSFR. Vol. 3. Bezobratlí. [Red book of endangered and rare plants and animals of Czechoslovakia. Vol. 3. Invertebrates]. Príroda, Bratislva, 155 pp (in Czech). TEYROVSKÝ V. 1930: Pøíspìvek k znalosti fauny vodních ploštic západní èásti Èeskoslovenské republiky. [Contribution to the knowledge of water bug fauna of the western part of the Czechoslovak Republic]. Sborník Klubu Pøírodovìdeckého v Brnì 12 (1929): 88–94 (in Czech, German summary without title). TEYROVSKÝ V. 1950: Pøedbìžný pøehled hlavních výsledkù výzkumu fauny vážek a vodních ploštic na Ostravsku a v oblasti Jeseníkù v srpnu (1.–6.), v záøí (2.–9.) a v øíjnu (27.–29.) 1950. [Preliminary overview of main results of the research on dragon-flies and water bugs in Ostrava and Jeseníky regions in August (1.–6.), September (2.–9.) and October (27.–29.) 1950. Pøírodovìdný Sborník Ostravského Kraje 11: 387–389 (in Czech). TROPEK R., KADLEC T., KARESOVA P., SPITZER L., KOCAREK P., MALENOVSKY I., BANAR P., TUF I. H., HEJDA M. & KONVICKA M. 2010: Spontaneous succession in limestone quarries as an effective restoration tool for endangered arthropods and plants. Journal of Applied Ecology 47: 139–147. TROPEK R., KADLEC T., HEJDA M., KOCAREK P., SKUHROVEC J., MALENOVSKY I., VODKA S., SPITZER L., BANAR P. & KONVICKA M. 2012: Technical reclamations are wasting the conservation potential of post-mining sites. A case study of black coal spoil dumps. Ecological Engineering 43: 13–18. VÁSÁRHELYI T. & BÁKONYI G. 2005: Typical aquatic and semiaquatic habitats and their Heteroptera fauna of the Lake Balaton. Folia Faunistica Hungarica 66: 39–49. VÁSÁRHELYI T., BÁKONYI G. & NOSEK J. 2005: A vízipoloska fauna évitzedes léptékû változása a Szigetközben. (Long-term changes in the aquatic Heteroptera fauna of the Szigetköz, North Hungary). Acta Biologica Debrecina, Supplementum Oecologia Hungarica 13: 249–258 (in Hungarian, English abstract). VÁSÁRHELYI T., NOSEK J., BÁKONYI G. & OERTEL N. 2007: Adatok a Ráckevei-(Soroksári-)Duna vízi és vízfelszíni-poloska valamint vízibogár faunájához (Heteroptera, Nepomorpha, Gerromorpha; Coleoptera: Hydradephaga, Hydrophiloidea). (Data to the aquatic Heteroptera and Coleoptera fauna of the Ráckevei- (Soroksári-)Danube, central Hungary (Heteroptera, Nepomorpha, Gerromorpha; Coleoptera: Hydradephaga, Hydrophiloidea)). Acta Biologica Debrecina, Supplementum Oecologia Hungarica 16: 221–229 (in Hungarian, English abstract). VINOKUROV N. N. 2009: Obzor vidov roda Teloleuca (Heteroptera, Saldidae) fauny Rossii i sopredel’nykh territoriy. (A review of species of the genus Teloleuca (Heteroptera, Saldidae) in the fauna of Russia and adjacent territories). Zoologicheskiy Zhurnal 88: 1189–1200 (in Russian, English summary) [English translation published in: Entomological Review, 2009, 89: 1092–1103]. VOTÝPKA J., KLEPETKOVÁ H., YURCHENKO V. Y., HORÁK A., LUKEŠ J. & MASLOV D. A. 2012: Cosmopolitan distribution of a trypanosomatid Leptomonas pyrrhocoris. Protist 163: 616–631. WACHMANN E., MELBER A. & DECKERT J. 2004: Wanzen. Band 2. Cimicomorpha. Microphysidae (Flechtenwanzen), Miridae (Weichwanzen). Die Tierwelt Deutschlands. 75. Teil. Goecke & Evers, Keltern, 288 pp. WACHMANN E., MELBER A. & DECKERT J. 2006: Wanzen. Band 1. Dipsocoromorpha, Nepomorpha, Gerromorpha, Leptopodomorpha, Cimicomorpha (Teil 1). Die Tierwelt Deutschlands. 77. Teil. Goecke & Evers, Keltern, 264 pp. WACHMANN E., MELBER A. & DECKERT J. 2007: Wanzen. Band 3. Pentatomomorpha I. Die Tierwelt Deutschlands und der angrezenden Meeresteile. Vol. 78. Goecke & Evers, Keltern, 272 pp. WACHMANN E., MELBER A. & DECKERT J. 2008: Wanzen. Band 4. Pentatomomorpha II. Die Tierwelt Deutschlands und der angrezenden Meeresteile. Vol. 81. Goecke & Evers, Keltern, 230 pp. WAGNER E. 1941: Conostethus hungaricus n. sp. (Hemipt. Heteropt. Miridae). Mitteilungen der Deutschen Entomologischen Gesellschaft 10: 76–80. WAGNER E. 1952: Blindwanzen oder Miriden. In: DAHL F. (ed.): Die Tierwelt Deutschlands, Bd. 41. Gustav Fischer Verlag, Jena, 218 pp. WAGNER E. 1965: Über einige bemerkenswerte Heteropteren aus dem Gebiet des Neusiedlersees. Wissenschaftliche Arbeiten aus dem Burgenland 32: 116–124. WAGNER E. 1966: Wanzen oder Heteropteren. I. Pentatomomorpha. In: DAHL F. (ed.): Die Tierwelt Deutschlands, Bd. 54. Gustav Fischer Verlag, Jena, 235 pp.

540 Acta Musei Moraviae, Sci. biol. (Brno), 98(2), 2013 Records of true bugs from Czech Republic and Slovakia V

WAGNER E. 1967: Wanzen oder Heteropteren II. Cimicomorpha. In: DAHL F. (ed.): Die Tierwelt Deutschlands und der angrezenden Meeresteile. Vol. 55. Gustav Fischer Verlag, Jena, 179 pp. WAGNER E. 1974: Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera, Heteroptera). Teil 2. Entomologische Abhandlungen Staatlichen Museum für Tierkunde Dresden 39 (Suppl.) (1973): 1–421. WAGNER E. & WEBER H. H. 1964: Hétéroptères Miridae. Faune de France. Vol. 67. Fédération Française des Sociétés de Sciences Naturelles, Paris, 590 pp. WIKARS L.-O. 1992: Skogsbränder och insekter. (Forest fires and insects). Entomologisk Tidskrift 113: 1–11 (in Swedish, English summary). WIKARS L.-O. 1997: Brandinsekter i Orsa Finnmark: biologi, utbredning och artbevarande. (Pyrophilous insects in Orsa Finnmark, Central Sweden: biology, distribution, and conservation). Entomologisk Tidskrift 118: 155–169 (in Swedish, English summary). WOLLMANN K. 1998: Corixidae (Heteroptera) in saueren Tagebauseen der Lausitz (Brandenburg). Lauterbornia 32: 17–24. WOLLMANN K. 2000: Corixidae (Hemiptera, Heteroptera) in acidic mining lakes with pH ≤ 3 in Lusatia, Germany. Hydrobiologa 433: 181–183. WRÓBLEWSKI A. 1980: Pluskwiaki (Hetemptera). Fauna s³odkowodna Polski. Vol. 8. Pañstwowe Wydawnictwo Naukowe, Warszawa – Poznañ, 156 pp (in Polish). ZÁHLAVOVÁ L. 2009: True bugs (Heteroptera) of the Žïár Nature Park, Czech Republic. Pp. 90–94. IN: SOLDÁN T., PAPÁÈEK M. & BOHÁÈ J. (eds.): Communications and Abstracts, SIEEC 21, June 28 – July 3, 2009. University of South Bohemia, Èeské Budìjovice, 96 pp. ZHAO Q., RÉDEI D. & BU W.-J. 2013: A revision of the genus Pinthaeus (Hemiptera: Heteroptera: Pentatomidae). Zootaxa 3636: 59–84.

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CONTENTS

EDITORIAL ...... 3

KMENT P., ŠTYS P. & GÜNTHER H.: On the occasion of the ninetieth birthday of RNDr. Jaroslav L. Stehlík, CSc...... 5

MALENOVSKÝ I., BURCKHARDT D. & DROSOPOULOS S.: RNDr. Pavel Lauterer, on the occasion of his eightieth birthday...... 11

OUVRARD D., BURCKHARDT D. & GREENWALT D.: The oldest jumping plant-louse (Hemiptera: Sternorrhyncha) with comments on the classification and nomenclature of the Palaeogene Psylloidea...... 21

BURCKHARDT D. & QUEIROZ D. L.: Phylogenetic relationships within the subfamily Aphalar- inae including a revision of Limataphalara (Hemiptera: Psylloidea: Aphalaridae). . . . . 35

WEBB M. D., RAMSAY A. J. & LEMAÎTRE V. A.: Revealing the identity of some early described European Cixiidae (Hemiptera: Auchenorrhyncha) – a case of ‘forensic’ taxonomy; two new combinations and a name change for Reptalus panzeri in Britain...... 57

OROSZ A.: A review of the genus Macrocixius with descriptions of five new species (Hemiptera: Fulgoromorpha: Cixiidae)...... 97

HOLZINGER W. E., HOLZINGER I. & EGGER J.: A new genus, Losirella, and two new species of Bennarellini from Ecuador (Hemiptera: Auchenorrhyncha: Fulgoromorpha: Cixiidae). 143

HOCH H., MÜHLETHALER R. & WESSEL A.: Acoustic communication in the subtroglophile planthopper Trigonocranus emmeae Fieber, 1876 (Hemiptera: Fulgoromorpha: Cixiidae: Oecleini)...... 155

ASCHE M.: The South-East Asian planthopper genus Lanaphora, with description of a new species (Hemiptera: Fulgoroidea: Delphacidae: Tropidocephalini)...... 163

GNEZDILOV V. M.: Notes on the genus Sarima (Hemiptera: Fulgoroidea: Issidae) with description of a new genus from Sri Lanka...... 175

MÜHLETHALER R. & GNEZDILOV V. M.: Two new species and additional records of the genus Eupteryx (Hemiptera: Cicadellidae: Typhlocybinae) from the northern Caucasus. . . . . 183

NIELSON M. W. & LOZADA P. W.: New species of Docalidia from Peru with new records (Hemiptera: Cicadomorpha: Cicadellidae: Coelidiinae)...... 191

D’URSO V., BÜCKLE C. & GUGLIELMINO A.: Platymetopius pavelitus sp.nov. (Hemiptera: Auchenorrhyncha: Cicadellidae), a new species from Italy...... 229

MALENOVSKÝ I.: New records of Auchenorrhyncha (Hemiptera) for the Czech Republic...... 235

MUSIK K. & TASZAKOWSKI A.: New data on some rare planthoppers and leafhoppers in Poland (Hemiptera: Auchenorrhyncha)...... 265

NICKEL H.: Zur Etymologie der Zikadennamen Mittel- und Nordeuropas. [On the etymology of Auchenorrhyncha names of central and northern Europe.] ...... 273 CONTENTS

BAÒAØ P. & Š TYS P.: Two new species of Oncylocotis (Hemiptera: Heteroptera: Enicocephalidae) from Australia...... 317

ZETTEL H.: Two new species of the enigmatic genus Aquulavelia (Hemiptera: Heteroptera: Veliidae) from Nepal and Bhutan...... 327

NIESER N., CHEN P.-P. & DE MELO A. L.: A new species and new synonymy in Limnocoris (Hemiptera: Heteroptera: Naucoridae) from Brazil...... 335

RÉDEI D.: Two new species of thread-legged assassin bugs from Australia (Hemiptera: Heteroptera: Reduviidae: Emesinae)...... 347

HEISS E. & BAÒAØ P.: Two new genera and four new species of micropterous Aradidae from Malaysia (Hemiptera: Heteroptera)...... 363

SCHAEFER C. W.: A new species of Dysdercus: Dysdercus stehliki sp.nov. (Hemiptera: Heteroptera: Pyrrhocoridae) from Brazil...... 381

DECKERT J.: Spilostethus stehliki sp.nov. (Hemiptera: Heteroptera: Lygaeidae) from Ethiopia...... 391

BRAILOVSKY H.: Description of four new species of Ninyas from Venezuela, a key to the known species and some new records (Hemiptera: Heteroptera: Geocoridae)...... 395

KONDOROSY E.: A new species of Kanadyana (Hemiptera: Heteroptera: Rhyparochromidae) from Ghana...... 407

KMENT P.: Carduelicoris stehliki, a new genus and species of Pentatomidae (Hemiptera: Heteroptera) from Madagascar...... 415

HANELOVÁ J. & VILÍMOVÁ J.: Behaviour of the central European Acanthosomatidae (Hemiptera: Heteroptera: Pentatomoidea) during oviposition and parental care...... 433

AUKEMA B., DUFFELS H., GÜNTHER H., RIEGER CH. & STRAUß G.: New data on the Heteroptera fauna of La Palma, Canary Islands (Insecta: Hemiptera)...... 459

KMENT P. H RADIL K., BAÒAØ P., BALVÍN O., CUNEV J., DITRICH T., JINDRA Z., ROHÁÈOVÁ M., STRAKA M. & SYCHRA J.: New and interesting records of true bugs (Hemiptera: Heteroptera) from the Czech Republic and Slovakia V...... 495