Progress in the Study of Sharpshooter Leafhoppers (Hemiptera: Auchenorrhyncha: Cicadellidae) Over 150 Years: Monographs, Museums and Individuals

Progress in the study of sharpshooter leafhoppers (Hemiptera: Auchenorrhyncha: Cicadellidae) over 150 years: monographs, museums and individuals

Michael R. Wilson & James A.Turner

The progress in taxonomic knowledge of the sharpshooter leafhoppers (Hemiptera: Cicadellidae: Cicadellinae) over the past 150 years is described. The availability of taxonomic monographs of the group has allowed an attempt to make digital images of the world fauna and to progress towards a web-based taxonomy. M.R. Wilson & J. A.Turner Department of Biodiversity and Systematic Biology, National Museum of Wales, Cardiff, CF10 3NP, UK. [email protected]

Introduction Abbreviations for Institutions discussed in text The leafhoppers comprise by far the largest family BMNH The Natural History Museum, London, within the Hemiptera, with approximately 19,500 UK described species in over 40 subfamilies (Oman et al. HNHM Hungarian Museum of Natural History, 1990) of which the subfamily Cicadellinae com- Budapest, Hungary prises around 2,400 species in around 330 genera. MMBC Brno, Czech Republic The name “sharpshooter” for this group of xylem- MNHN Muséum National d’Histoire Naturelle, feeding leafhoppers has increasingly been used espe- Paris, France cially in the USA. They are among the largest and NCSU North Carolina State University Insect most brightly coloured of the leafhoppers. Some spe- Collection, Raleigh, North Carolina, USA cies are important vectors of the xylem-limited bacte- NHRS Naturhistoriska riksmuseet, Stockholm, rium Xylella fastidiosa, which affects both citrus trees Sweden in Brazil as well as grapevines in southern USA. NMW Naturhistorisches Museum Wien, Vienna, The Cicadellinae, as currently defined, was revised Austria by David Young (1915–1991) in three remarkable USNM National Museum of Natural History, [for- volumes (Young 1968a, 1977a, 1986a). The pub- merly, United States National Museum], lication of these works has enabled the evaluation Washington, DC, USA and description of additional genera and species, ZMHB Museum für Naturkunde der Humboldt- primarily by researchers in Brazil and China. The Universität, Berlin, Germany availability of these taxonomic monographs and subsequent publications has made the Cicadellinae a relatively well-known group. In 2004 a project Taxonomic history was started, funded by UK-based Leverhulme Trust, which has enabled a compilation of digital images of A short history of cicadelline study Cicadellinae. This paper is a brief review of the project in the con- Linnaeus (1758a) described four cicadelline spe- text of the history of taxonomic work on this group cies in the genus Cicada, which included many of leafhoppers over the past 150 years. other Auchenorrhyncha species, such as treehoppers,

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2500

2000

Young

1500

Melichar Fowler / 1000 Distant

Walker / Signoret 500

0 1750 1775 1800 1825 1850 1875 1900 1925 1950 1975 2000 Fig. 1. Species accumulation curve from 1750 to present, based on valid species.

spittlebugs, planthoppers, and true cicadas. A first erected himself, he treated all species in the large ge- attempt to better define the genus Cicada was nus Tettigonia. He regretfully decided to abandon the made by Fabricius (1794a), who transferred some previously established genera because he understood Linnaean species of other Auchenorrhyncha and their diagnostic characters as part of a continuum, even some leafhoppers (such as Jassus) to other gen- impossible to objectively break down, event though era. Following this Germar (1833a) defined the he did assemble Tettigonia species in five groups. genus Tetigonia Geoffroy (posteriorly emended to Stål (1869) attempted to separate genera and de- Tettigonia by Olivier, 1789a, but rejected by the scribed other new genera in the subfamily. His ar- International Commission of Zoological Nomen- rangement was mostly followed by Fowler in a se- clature, see Hemming 1954, and currently used as ries of papers (1894–1909) in the “Biologia Centrali synonymous to Cicadella Latreille, ICZN 1963a) us- Americana”. This volume was also accompanied by ing characters that will mostly define the subfamily high quality colour illustrations. Fowler treated a Cicadellinae for following authors, such as the tumid number of genera that would all fall into the tribe frons and the ocelli positioned on the middle of the Proconiini as presently defined. His remaining spe- crown. Since Linnaeus (1758) about 2,790 species cies were placed in Tettigonia and he covered 160 have been described in the tribes Cicadellini and species (in what would be the tribe Cicadellini). Proconiini by around 60 authors. Of the described Fowler (1899: 235) also states that: “It is possible species, around 2,290 are currently accepted as valid. that the genus may at some future time be divided, Signoret (1853b,c, 1854a,b, 1855a,b,c,d) was the but it seems impossible to do this satisfactorily in first to attempt to monograph the group as it was the present stage of our knowledge”. As with most then understood. He published re-descriptions of a authors at this period, Fowler was relying on exter- number of species (including those of Walker, with nal characteristics, as did also Distant (1908, 1918) whom he was a contemporary) and described a large who also described many species from the Oriental number of species, accompanied by colour illustra- region. tions (e.g. see Fig. 2). Although Signoret (1853) rec- Melichar (1924a, 1925a, 1926a, 1932a, 1951a) ognized that there were twelve generic names availa- made a substantial contribution in his monograph- ble, including Dilobopterus Signoret, 1850, which he published after his death in 1924. He divided the

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Fig. 2. Reproduction of one of the Plates from the Signoret monograph. – 1, Erythrogonia colorata (Germar, 1821); 2, ‘Tettigonia’ divisa Signoret, 1853; 3, Tettisama quinquemaculata (Germar, 1821); 4, Erythrogonia areolata (Signoret, 1853); 5, Erythrogonia jucunda (Walker, 1851); 6, Erythrogonia quadriguttata (Fabricius, 1787); 7, Erythrogonia sexguttata (Fabricius, 1803); 8, Erythrogonia quadriplagiata (Walker, 1851); 9, Trachygonalia germari (Signoret, 1853); 10, Cardioscarta quadrifasciata (Linnaeus, 1758); 11, Agrosoma pulchella (Guerin- Meneville, 1829); 12, Agrosoma proxima (Signoret, 1853); 13, Agrosoma cruciata (Signoret, 1853); 14, Stehlikiana crassa (Walker, 1851) = walkeri Signoret, 1853; 15, Graphocephala multicolor (Signoret, 1853); 16, Tettigoniella cosmopolita (Signoret, 1853).

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subfamily into two sections: the Proconiaria with 54 genera and the Cicadellaria with 101 genera, which correspond more or less with the Cicadellini and Proconiini as treated subsequently by Young. The main problem with using Melichar’s classification is that the work is entirely without illustrations. While further species were described by other workers following Melichar, it was not until Young commenced his work in around 1952 that real progress was made, although the availability of the Metcalf catalogue to the “Tettigellinae” (1964) would have made progress easier. In the introduction to his monograph on Proconiini (Young 1968a) he states that the need for a generic revision became clear while working for the US Department of Agricul- ture. Cicadellinae specimens sent for identification could rarely be identified to genus on the basis of existing literature. Young followed previous authors (e.g., Oman 1949) by utilising characters of the male genitalia to delimiting genera (in addition to using female terminalia as specific diagnostic characters in 1977a and 1986a) and a move to the University of North Carolina provided more research time than available before. His monographic revisions of the group (Young 1968a, 1977a, 1986a) occupied him for the rest of his career, and has given a remarkable legacy from which all modern work has started. Fig. 3. Victor Antoine Signoret (taken 1865). From While Young’s intention was to provide a generic DEI, Muncheburg, Germany (Groll & Schubert framework, he described many new species and also 2006). made synonyms based on his studies in European museums. He did not, however, provide a modern phylogenetic analysis of the relationships of the gen- Signoret, Victor Antoine (1816–1889) (Fig. 3) era, and only gave schematic relationships among Victor Signoret was born in Paris, France April 6 genera and groups of genera. The availability of the 1816 and died in Paris on April 3 1889. Signoret monographs has allowed a new generation of special- was a qualified pharmacist and medical doctor and ists, especially in Brazil and China, to describe many practised in Paris. He started his natural history new species. Recently some re-analysis of generic studies with Coleoptera, but moved his interests and species relationships using phylogenetic analysis to Hemiptera and published around 130 scientific both morphological and molecular data and has been papers mostly on Hemiptera. He was a founder of carried out in the tribe Proconiini (Takiya 2007). scale insect taxonomy and made slide preparations A new catalogue to the Cicadellinae has been pro- of specimens in order to study their morphology duced (McKamey 2007) which will make the group (Ben-Dov & Matile-Ferrero 1995). These authors further accessible. also indicated that notebooks from his later studies were stored in the MNHN. These show Signoret to be a superb artist and his pencil sketches of Heter- Who described the species optera (especially Cydnidae) and leafhoppers are As discussed above, around 60 individuals have de- quite remarkable. Among the major achievements scribed species in the Cicadellinae since Linnaeus of his studies on Auchenorrhyncha is the “Revue (1758). The accumulation of species is shown (for icongraphique des Tettigonides”, published in the valid species) in Fig. 1. But around 65% of the to- “Annales de la Société Entomologique de France” in tal currently valid species (2,300 species) have been parts between 1853–1855. The final part contains described by just six individuals (indicated on the an index to species. Figure). It is worth presenting their work in more Signoret reviewed the known sharpshooter species detail. and described 246 species (of which 211 are presently considered valid). The highlight of the work

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Fig. 4. Francis Walker (coll Kraatz) From DEI, Munche- Fig. 5. William Lucas Distant. From BMNH Entomol- burg, Germany (Groll & Schubert 2006). ogy Library, London, UK.

is the 19 beautifully illustrated colour plates of the scribing the same species more than once under dif- dorsal view of 310 species (e.g. Fig. 2). It is inter- ferent specific names. The British Museum paid him esting that, as noted by Young (1968a), the colour one shilling for each new species and one pound for of the plates varies between different copies. All spe- each new genus. He is best known for his catalogues cies were treated in the genus Tettigonia and Signoret of Orthoptera, Neuroptera, Homoptera, Diptera, indicated in the text the collection from which the Lepidoptera and Hymenoptera. Upon his death specimens originated. one (anonymous) obituary read: “More than twenty Upon his death in 1889, his collection of around years too late for his scientific reputation, and after 30,000 specimens was sold to the NMW. There having done an amount of injury to entomology al- were 3000 species of Auchenorrhyncha represented most inconceivable in its immensity, Francis Walker by 8,300 specimens and among these were many of has passed from among us.” Walker described 222 the specimens used in the preparation of the mono- Cicadellinae species of which 138 are considered graph. The NMW collection was studied by Young valid. Despite his generally poor taxonomic reputa- in 1962–63 and he designated lectotypes for most tion, Carvalho & Webb (2005) describe in detail the species (Young & Beier, 1963a). Significant numbers labelling of Walker specimens, which has generally of Signoret specimens are also found in the ZMHB. led to confident selection of type specimens of the species he described. Walker, Francis (1808–1874) (Fig. 4) Francis Walker was born in Southgate, England on Distant, William Lucas (1845–1922) (Fig. 5) July 1 1809 and died at Wanstead, England October William Distant was born in London on 12 Novem- 5 1874. Walker was employed by the BMNH as a ber 1845 and died in London on 4 February 1922. curator between 1844 and 1873. He described al- He had a keen interest in natural history from an most 20,000 new insect species, but, unfortunately, early age and he developed an interest in the taxon- he was, sometimes a careless taxonomist, often de- omy of the Coleoptera, Lepidoptera and Hemiptera.

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Fig. 6. William W. Fowler. From Entomologist’s Fig. 7. Leopold Melichar. From MMBC, Czech Monthly Magazine 1923 Vol 59: facing page 152. used Republic. with permission of the Journal.

In his later life he concentrated almost completely his degree at Oxford William Fowler was ordained on the Hemiptera and especially the Cicadidae. He a priest and eventually appointed as Canon in Lin- was appointed as a part-time assistant at the BMNH, coln Cathedral. He was drawn in his leisure time to London in April 1899, where he worked on the ar- the study of insects and especially Coleoptera and rangement of the Hemiptera for two or three days wrote “The Coleoptera of the British Islands” which a week until ill health made it impossible for him appeared in five volumes between 1887 and 1891. to continue. He published the results of his stud- Despite its age this book is still highly sought after ies at the Museum in a series of papers “Rhyncho- for the study of Coleoptera in Britain. His inter- tal Notes” in the “Annals and Magazine of Natural ests were not confined to British entomology and History”. He also contributed to Volume 1 of the he wrote the Hemiptera-Homoptera sections of the Homoptera of the Biologia Centrali-Americana and “Biologia Centrali-Americana”, which appeared the Hemiptera volumes of the Fauna of British India. between 1894–1909. This work also contains Distant published several hundred titles and many of some beautiful coloured plates (e.g Fig. 9) (which them were divided into parts. His bibliography was are available online at http://www.sil.si.edu/ compiled by Dolling (1991). digitalcollections/bca/). He described 139 (113 valid) Distant described 147 cicadelline species (79 Old Cicadellinae species and the majority of these are World and 68 New World) of which 104 species are housed in London, but some others are in Vienna currently accepted. The majority of the type material (Wilson 2008). Young (1965c) selected and desig- is deposited in the BMNH, London. nated lectotypes of Fowler’s species in the course of preparation of his monographs. Fowler, William W. (1849–1923) (Figs 6, 9) William Fowler was born in Gloucester, England in Melichar, Leopold (1856–1924) (Fig. 7) January 1849 and died in Reading on June 3, 1923. Leopold Melichar was born in Brno, Moravia, Canon Fowler’s father was a Vicar and after taking Czech Republic on December 5 1856 and died in

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morphology, and the majority of his new genera, although validly published according to the no- menclatural rules of the International Commission of Zoological Nomenclature (ICZN 1999), lacked formal descriptions and type designations. China (1927d) dealt with preoccupied generic names from Melichar (1926a) and later (in 1938d) designated type species for several of Melichar’s genera based on the remaining unpublished manuscript. During his European study visit in 1962–63 David Young spent some weeks in Brno and fully examined the Melichar collection and subsequently designated lectotypes for Melichar species deposited in the MMBC (Young & Lauterer 1966a). Further Meli- char lectotype designations were made by Lauterer & Schroder (1970). Young also worked extensively on the specimens and the Cicadellinae collection is now fully re-curated by Pavel Lauterer and is the best represented after Raleigh, London and Washington. A visit to HNMH, Budapest in 2006 enabled MRW to study the cicadelline collection and inventory the Melichar species in that museum (Wilson & Takiya 2007).

Young, David Allan (1915–1991) (Fig. 8) David Young was born May 26 1915, Wilkinsburg, Fig. 8. David Allan Young. From NCSU Archive, used Pennsylvania, USA and died June 8 1991, Kentucky. with permission. He studied entomology at Cornell University and received his M.S. in 1942. Following war service, he took an instructorship in the Department of Biology at the University of Louisville until 1948, Brno on September 2 1924. For most of his career after which he entered the University of Kansas to Melichar was a medical doctor in Vienna. In 1912 study for his Ph.D., awarded in 1950, under the he retired to Brno to allow more time to be spent on direction of R. H. Beamer. From 1950 to 1957 he entomology. After his death his insect collection was worked at the USNM employed by the US Depart- passed to the MMBC where it is currently housed. ment of Agriculture. In 1957, Young accepted a Melichar was the first since Victor Signoret to at- position as an associate professor at North Carolina tempt a monographic treatment of the Cicadellinae, State University, where he continued his systematic but no part was published during his life. As well research on leafhoppers (begun with Beamer) and as his own collection, he also examined specimens administered the NCSU Insect Collection. Young from other European museums, especially from (1957) published a synopsis of the U.S. species of HNHM. His manuscript was sent to Dr Géza Hor- Homalodisca (Proconiini) and, after his promotion to váth for publication in the Annales Musei Nationales Professor in 1961, spent the academic year in Europe Hungarici where four parts appeared between 1924 (1962–1963) visiting a number of European muse- and 1932. However, in 1951, following questions ums where the large majority of species, which had about the completion of the series and the remain- been described by European workers were deposited. ing manuscript, Dr Vilmos Székessy published a This visit enabled Young to study most of the types further part in Melichar’s name (Melichar 1951a). of Cicadellinae described by European workers and In these monographs, seventy-seven new cicadel- the taxonomic and nomenclatorial research became line genera and approximately 200 taxa in the spe- the basis of the three monographs he produced on cies-group were described. These species-group taxa the group (Young 1968a, 1977a & 1986a).). We include many “varieties”, which were correctly in- now take it for granted that we take a laptop compu- terpreted as available names of subspecific rank by ter with us to visit museums, in order to take notes Young. Unfortunately, the work was entirely without on specimens examined, perhaps we may also have a illustrations, characters were based solely on external database and illustrations to assist our studies. None

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of this was available to Young during his year-long was any contact between these authors and Melichar. visit. Young published a series of papers designat- After Melichar’s death in 1924, there was a gap of ing lectotypes based on these studies: Stockholm, almost 35 years before David Young started his work Sweden (mostly Stål species) (Young 1963a); Berlin, on these insects. Signoret, Melichar, and Young all Germany (Young 1964a); Eberswalde, Germany intended a monographic treatment but Young de- (Breddin species) (Young 1965a); London, U.K. scribed many more species than the other workers (Walker, Distant and Fowler species) (Young 1965c); combined. This was mostly, of course, because he Copenhagen, Denmark and Kiel, Germany (Fabricius was the first to use the male genitalia structures in species) (Young 1965d); Dresden, Germany (Jacobi the Cicadellinae in any detail. He was also able to species) (Young & Lauterer, 1964a); Brno, Czech Re- both examine most of the type material and accumu- public (Melichar species) (Young & Lauterer 1966a); late large collections for study. Warsaw, Poland (Schmidt species) (Young & Nast Signoret and Melichar were both medical doctors 1963a); Budapest, Hungary (Young & Soós 1964a); but their remarkable entomological outputs would Paris, France (Young 1974). The political situation impress many “professionals”. Walker was, of course, in parts of Europe made it difficult or impossible a generalist who described species in many groups. to visit certain museums. There were a number of Distant and Fowler worked on other groups as well cicadelline species (around 150 species) that Young as Hemiptera. Young was the only one of the 6 who either failed to place in a genus (especially unique concentrated almost exclusively on the Cicadellinae female types) or failed to locate during his studies. and was a professional taxonomist (and one who He listed these at the end of each of the monographs trained others by supervising undergraduate and (Young 1977a, 1986a) as genera and species of un- graduate studies). certain position. The format of Young’s monographs was established in the 1968 study of the tribe Proconiini. In general Where are the species deposited? lectotype designations had been made prior to pub- The accumulation of specimens in any one museum lication, but frequently, taxonomic notes on these is a result of geography, history, staff, and collectors. were given in the monographs. Unfortunately, for Despite the majority of species being found in the reason of time constraints, Young did not list muse- tropics of almost all of the world, Cicadellinae spe- ums where he examined specimens of previously de- cies described until the 1950’s had been by European scribed species, or provide detailed descriptions, but workers. Much of this material had been deposited did usually figure the characters of the male genitalia in major museums in central and northern Eu- (often of a lectotype or topotype). Full details and rope. During the study visit made by David Young descriptions were only provided for new species. A in 1962–63 much of this was examined, lectotypes list of species with synonyms and country records were designated, and he also loaned large numbers of is given for each genus. Effectively each genus dis- unidentified specimens. As discussed above, Young cussed is a partial revision and in some cases it is did not give the details of the museums from where now clear that further work is necessary to provide he studied specimens of the existing species. Dur- a full revision. Among all three monographs, Young ing the current imaging project (see below) we have described 738 species (734 currently valid) among accumulated information on over 30 institutions, 169 genera. which house significant collections in this group. On completion of his monographs the specimens Interactions between workers were returned to their original museums, but sev- The major workers have all followed and built on eral representatives were retained for deposition in each other’s work in the 150 years since the publica- the NCSU collection, Raleigh. As a result, Raleigh tion of the first part of Signoret’s monograph. He has the largest number of species represented, fol- had synthesised the earlier accounts in the group, but lowed closely by the BMNH, London and the few other workers were specialised in leafhoppers. USNM, Washington, D. C. Each of these institu- There is evidence that Signoret had been in contact tions has around 50% of the world fauna represent- with Walker and indeed examined some of his speci- ed. The BMNH is notable for both the number of mens during a visit to London. Both Signoret and Cicadellinae species and of historic types (Walker, Walker had died before Melichar commenced his Distant, and Fowler). monographic treatment. Fowler and Distant were Other large museums have between 350 (NMW, contemporaries, but cryptic remarks in the introduc- HNHM) and 560 (NHRS) species but are very tion to the Biologia Centrali-Americana suggest lit- important because of the number of historic types tle collaboration. There is little evidence that there present. The Bishop Museum, Honolulu has 75% of

Downloaded from Brill.com10/03/2021 04:41:22AM via free access Wilson & Turner: Sharpshooter leafhoppers over 150 years 297 the species known from New Guinea (110 species: Leverhulme Trust in 2004. The intention was to take almost all endemic), although representatives are images that would provide a web-accessible informa- found in London (36%) and in Raleigh (51%). Ta- tion source, as well as volumes of images that would ble 2 gives information on the numbers of cicadelline reflect the three monographs by Young. species described by the 6 main authors and their presence in each of the 5 main collections. Perhaps Methods not surprisingly the majority of species are found in Each photograph was made using JVC KY70 3CCD the “home” Institute (or place where the collection is camera on a Leica microscope MZ8 and Synoptics deposited) of the author. This effect is especially seen Automontage software. This software allows for a in Fowler and Distant species, which are overwhelm- number of images (we usually used around 20) each ingly represented in the BMNH, London. This may at a slightly different focus to be “montaged” to pro- also be an effect of their working on a regional fauna. vide an image with an enhanced depth of field. Im- It is less obvious among Signoret species, perhaps ages were saved both as TIFF files and as JPG (for because he was an early pioneer whose species are use in the database). Each TIFF image was around likely to be among the more commonly encountered 4MB in size. A special chamber (designed by James and also because he described on a world basis. The Turner, who made almost all of the images) was used NCSU, Raleigh also has the largest percentage of diffusing the lighting in order to reduce shadows and species described by David Young, although com- highlights created by intense fibre optic light. pared to other authors, Young species are represented in a lower percentage (see Table 2). Young did retain Whenever possible, specimens were selected that some representatives in NCSU when he described had been card pointed so that no pin had been used. species, but he also studied specimens from many Where pins had been used for mounting they were other institutions. Additionally, Young described “removed” using Adobe Photoshop, as were the legs, many species based on a single specimen and he did so that each specimen was consistent. Minor damage deposit all his primary types in the USNM. to specimens was also “repaired” using Photoshop. However, there would be no doubt as to which speci- Interestingly only around 20 cicadelline species are men has been illustrated, because information as to found in all of the ten largest collections. Even the the museum where the specimen is stored is given top seven contain only 86 species, but this number with the image. All images were managed in the rela- rises to 170 in the largest five institutions, and to tional database FilemakerPro v.8.5, comprising taxo- 281 in the top four. The overlap in the three largest nomic, image and depository databases. Images are institutions for this group is, however, considerable, given a unique reference number, which has been at- and 468 species are found in all of these, and be- tached to the imaged specimen as a label. Museums tween NCSU and BMNH, 694 species are found in from which imaged specimens have been loaned will common. Naturally, in common with most insects, be provided with a spreadsheet with this information the majority of cicadelline species are known only and copies of the images. from the type material, with 282 species found only in the largest five collections (with BMNH having Choice of specimens 120 of these). Most institutions have numerous spe- Some have assumed that during this project type cies unique to them. material would be imaged as a priority. But many historic type specimens are in relatively poor condi- tion and unsuitable to provide good quality images. The current project Given that David Young examined many of these The current project to image Cicadellinae species on type specimens the selection of specimens to pho- a world basis arose by having seen a large collection of tograph was initially based on the quality of avail- unidentified sharpshooters in Quito, Ecuador. Many able specimens identified by him in the course of his of these could be identified to genus and to species work. Following the publication of the monographs if access to a good collection and the monographs by other specialists have identified some species. Many Young, were possible. Neither of these were available species, assumed to be those that are more common in Ecuador. Another approach was to accumulate im- and widespread, are to be found in a number of ages of species that might be placed on a website or museums and there was plenty of choice of suitable in a modern monographic treatment, which would specimens. Many species are known from only small nevertheless be based on the work of David Young numbers of specimens in the type series located in and subsequent studies. It was this approach that re- a limited range of museums. In practice, each mu- sulted in a successful application for funding by The seum visited or contacted during the course of this

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Fig. 9. Reproduction of a plate from Fowler, Biologia Centrali-Americana.

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Fig. 10. Selection of images from current project. – 1, Agrosoma proxima (Signoret, 1853); 2, Agrosoma pulchella (Guérin-Méneville, 1829); 3, Cardioscarta quadrifasciata (Linnaeus); 4, Erythrogonia areolata (Signoret, 1853); 5, Erythrogonia colorata (Germar, 1821); 6, Erythrogonia jucunda (Walker, 1851); 7, Erythrogonia quadriplagiata Walker, 1851; 8, Tettisama quinquemaculata (Germar, 1821); 9, Graphocephala multicolor (Signoret); 10, Stehlikiana crassa (Walker, 1851); 11, Tettigoniella cosmopolita (Signoret, 1853); 12, ‘Tettigonia’ divisa Signoret, 1853; 13, Oncometopia (Similitopia) alpha (Fowler, 1899); 14, Phera lacerta Fowler, 1899; 15, Oncometopia undata (Fabricius, 1794).

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Table 1. Number of species in five largest Cicadellinae collections. Number of valid species: 2281 (Old World 640; New World 1641)

Institute Total No. No. & % Old World No. & % New World No of species (%) species species ‘uniques’

NCSU, Raleigh 1121 (49) 228 (36) 893 (54) 18 BMNH, London 1030 (45) 286 (44) 744 (45) 120 USNM, Washington 954 (42) 149 (23) 805 (49) 55 MMBC, Brno. 715 (32) 126 (20) 589 (35) 43 NHRS, Stockholm 563 (28) 99 (15) 464 (28) 30

Table 2. Number of species (and % of total) from each of main authors in the five largest Cicadellinae collections.

Signoret Walker Fowler Distant Melichar Young 210 138 113 104 165 745

NCSU, Raleigh 115 (55) 74 (54) 72 (64) 51 (49) 64 (39) 425 (57) BMNH, London 109 (52) 123 (90) 107 (95) 100 (96) 39 (24) 261 (35) USNM, Washington 93 (44) 52 (38) 58 (51) 30 (29) 62 (38) 383 (51) MMBC, Brno 97 (46) 58 (42) 35 (31) 41 (39) 131 (79) 141 (19) NHRS, Stockholm 103 (49) 57 (41) 21 (19) 31 (30) 39 (24) 148 (20) NMW, Vienna 146 (70) - - - 55 (33) -

work enabled a new range of species to be added to Young’s list at the end of his 1977 and 1986 mono- the database. Also, Young described 745 species so graphs. One unexpected and satisfying outcome of the specimens he utilised in the description of each the project and especially of having been able to visit species may be regarded as authorative. a number of important European museums, was being able to locate types of around 30 species that Time constraints did not allow us to take more than Young failed to see during his studies. the dorsal view of each species. However, for the Proconiini a lateral view was also taken because the The Future lateral view head shape is often diagnostic. A view of Since the compilation of Young’s monographs, the face would also have been very useful for species there has been a rapid acceleration of taxonomic recognition in some genera. A number of specimens work being published on the group, especially by were imaged if sexual dimorphism, colour, or pattern workers in Brazil and China. Most of these works variation was noted during study. involve descriptions of new species and small revisions. A morphological character and molecular Project Outcomes analysis on the tribe Proconiini has been carried During the two years of the Leverhulme project out (Takiya 2007), from which the morphologi- around 5,000 images have been taken and 90% of cal part has generated an online interactive key to the world Cicadellinae fauna represented at species the genera and taxonomic database including geo- level (e.g Fig. 10). Over 95% of genera have been graphical, host-plant, images, and literature data imaged. A website has been completed, based on the (http://ctap.inhs.uiuc.edu/takiya). It is anticipated FileMakerPro database. Three volumes of images that the availability of Young’s monographs, the new will be produced as an illustrated checklist (based on taxonomic catalogue (McKamey 2007), and online McKamey 2007), with the first being a volume on accessible interactive identification tools, such as in- Old World Cicadellini (2007 in prep.). In this vol- teractive keys and images resulting from this project, ume, for ease of use, the species have been divided will enhance the rate of further revisionary work in into sections relating to geographical regions in the this group. Old World. There is little overlap between species. The section “Species of uncertain position” reflects

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Acknowledgements emendata et aucta. Secundum classes, ordines, genera, This project would not have been possible were it species adjectis synonimis, locis, observationibus, de- not for the grant from The Leverhulme Trust, which scriptionibus 4: 1–472. has funded the employment of the co-author as Re- Fowler W.W. & T.D.A. Cockerell., 1894–1909. Insecta search Assistant to MRW at the National Museum Rhynchota. Hemiptera-Homoptera. Volume II, of Wales, Cardiff. This funding also allowed visits to Part 1. [Available ��online at http://www.sil.si.edu/ digitalcollections/bca/)]. BMNH, London, UK; USNM, Washington DC, Groll, E.K. & Schubert, E. 2006. Digitale Entomologische USA and Museu Nacional, Rio di Janiero, Brazil. Information - Images. Database 1. version, DEI im Additional funding for visits to Hungarian Natural ZALF e.V. http://www.zalf.de/home_zalf/institute/ History Museum, Budapest, Hungary; Muséum dei/php_e/archiv/bilder.php National d’Histoire Naturelle, Paris, France; Polish Hemming, F. 1954. Opinion 2999. Validation, under Academy of Science, Museum of the Institute of plenary powers, of the generic names Tettigonia and Zoology, Warsaw, Poland; Naturhistoriska Acrida, in the Order Orthoptera (Class Insecta) riksmuseet, Stockholm, Sweden; Naturhistorisches as from Linnaeus, 1758 (Ruling supplementary Museum Wien, Vienna, Austria; and Museum für to the ruling given in opinion 124). International Naturkunde der Humboldt-Universität, Berlin, Commission Zoological Nomenclature – 8: 209–235. Germany was provided from the EU Synthesys [ICZN] International Commission of Zoological Nomen- Programme. We are grateful to Dr Stuart McKamey clature, 1963a. Opinion 647. Cicadella Latreille, 1817 and Dr Daniela Takiya for reading the ms and (Insecta: Hemiptera); validation under the plenary helpful comments. powers. – Bulletin of Zoological Nomenclature 20: 35–38. The Entomology Library BMNH kindly provided [ICZN] International Commission of Zoological No- the portrait of W.L. Distant. menclature, 1999. International Code of Zoological Igor Malenovsky is thanked for his hospitality while Nomenclature. – London: International Trust for Zo- visiting MMBC and for providing the portrait of ological Nomenclature. Fourth Edition. L. Melichar. Lauterer, P. & H. Schröder, 1970a. Types of Cicadellinae (Homoptera, Cicadellidae) in the Moravian Museum. – Casopis Moravskeho Musea v Brno 55: 127–132. References Linnaeus, C., 1758a. Hemiptera. 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Agriculture Research Service 730 pp. Walker, F., 1858a. Homoptera. Insecta Saundersiana: or Olivier G.A.,1789a Hémiptères, section 1. – Encyclopédie characters of undescribed insects in the collection of méthodique histoire naturelle insects. 4 1–331. William Wilson Saunders, Esq. – 117 pp. Oman P.W., 1949 The Nearctic leafhoppers (Homoptera: Walker, F., 1858b. Supplement. List of the specimens of Cicadellidae) A generic classification and check list. Homopterous insects in the collection of the British – Memoirs of the Entomological Society of Washing- Museum. – 307 pp. ton 3: 1–353. Walker, F., 1870b. Catalogue of the Homopterous Oman, P.W., Knight, W.J. & M.W. Nielson, 1990a. Insects collected in the Indian Archipelago by Leafhoppers (Cicadellidae): a Bibliography, Generic Mr A.R. Wallace, with descriptions of new species. Check-list and Index to the World Literature 1956– – Journal Proceedings Linnean Society London 10: 1985. – CAB International Institute of Entomology, 276–330. Wallingford, UK. [iv] + 368 pp. Wilson, M.R. & D.M. Takiya., 2007. Cicadellinae (Hemi- Signoret V.,1850b. Description d’un genre nouveau et de ptera, Auchenorrhyncha: Cicadellidae) described by quelques espèces du groupe des tetttigonides. Revue Leopold Melichar in the Hungarian Natural His- et Magazin de Zoologie Oure et Appliquée. (2) 2: tory Museum. – Annales Historico-Naturales Musei 283–289. Nationalis Hungarici 99: 1–22. Signoret, V., 1853b. Revue iconographique des Tet- Young, D.A., 1958a. A synopsis of the species of Homalo- tigonides. – Annales de la Société Entomologique de disca in the United States. – Bulletin of the Brooklyn France 1 (3) 323–374, pls 8–12. Entomological Society 53(1): 7–13. Signoret, V., 1853c. Revue iconographique des Tet- Young, D.A., 1963a. Types of Cicadellinae (Homoptera, tigonides. – Annales de la Société Entomologique de Cicadellidae) in the Naturhistoriska Riksmuseet in France 1 (3) 661–688, pls 21–22. Stockholm. – Entomologisk Tidskrift 84: 224–227. Signoret, V., 1854a. Revue iconographique des Tet- Young, D.A., 1964a. Some cicadelline types of species de- tigonides. – Annales de la Société Entomologique de scribed by Signoret from Berlin collections (Homop- France 2 (3) 5–28 pls 1–2. 341–366 pls 11–12, 483– tera Cicadellidae). – Mitteilungen aus dem Zoolo- 496 pl 17, 717–732 pl 21. gischen Museum in Berlin 40: 9–13. Signoret, V., 1854b. Revue iconographique des Tet- Young, D.A. 1965b., Some cicadelline lectotypes of Bred- tigonides. – Annales de la Société Entomologique de din species. – Beitraege zur Entomologie 15(1/2): France 2 (3) 341–366, pls 11–12. 11–14. Signoret, V., 1855a. Revue iconographique des Tet- Young, D.A., 1965c. Cicadelline types in the British Mu- tigonides. – Annales de la Société Entomologique de seum (Natural History) (Homoptera: Cicadellidae). France 3(3) 49–60, pl 6. – Bulletin of the British Museum (Natural History) Signoret, V., 1855b. Revue iconographique des Tet- Entomology 17(4): 163–199. tigonides. – Annales de la Société Entomologique de Young, D.A., 1965d. Notes on Fabrician species of France 3(3) 225–240 pls 6,12. Cicadellinae (Homoptera, Cicadellidae) in Copenha- Signoret, V., 1855c. Revue iconographique des Tet- gen and Kiel with lectotype designations. – Entomolo- tigonides. – Annales de la Société Entomologique de giske Meddelelser 34(1): 10–18. France 3(3) 507–528 pl 21. Young, D.A., 1968a. Taxonomic study of the Cicadellinae Signoret, V., 1855d. Revue iconographique des Tet- (Homoptera, Cicadellidae), Part 1. Proconiini. – Tech- tigonides. – Annales de la Societé Entomologique de nical Bulletin of the United States National Museum France 3(3) 765–836 pls 23–24. 261: 1–287. Stål, C., 1869. Hemiptera Fabriciana 2 – Svenska Vetensk Young, D.A., 1974a. Types of Cicadellinae (Homoptera, Akad Handl. 8(1) 231–300. Cicadellidae) in the Museum National d‘Histoire Takiya, D.M., 2007. Systematic studies on the leafhop- Naturelle, Paris, France. – Bulletin Museum Na- per subfamily Cicadellinae (Hemiptera: Cicadellidae). tional d‘Histoire Naturelle, Serie III (Paris) 263: Unpublished PhD thesis. – University of Illinois at 1697–1699. Urbana-Champaign.Pp. 183. Young, D.A., 1977a. Taxonomic study of the Cicadel- Walker, F., 1851b. List of the specimens of Homopter- linae (Homoptera: Cicadellidae), Part 2, New World ous insects in the collection of the British Museum. Cicadellini and the genus Cicadella. – Technical – British Museum, London. 3: 637–907. Bulletin of the North Carolina Agricultural Experi- Walker, F., 1857a. Catalogue of the Homopterous ment Station 239: 1–113. Insects collected at Singapore and Malacca by Young, D.A., 1979a. A review of the leafhopper genus Cofa- Mr A.R. 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(Homoptera, Cicadellidae) in the Hungarian Natural Young, D.A. & P. Lauterer., 1964a. Cicadelline lectotypes History Museum. – Annales Historico-Naturales from the A. Jacobi collection (Homoptera, Cicadelli- Musei Nationalis Hungarici 56: 465–467. dae). – Reichenbachia 2: 293–296. Young, D.A. & J. Nast., 1963a. Cicadelline types of spe- Young, D.A. & P. Lauterer., 1966a. Types of Cicadellinae cies described by Edmund Schmidt (Homoptera, (Homoptera, Cicadellidae) in the Moravian Museum. Cicadellidae). – Annales Zoologici (Warsaw) 21: – Casopis Moravskeho Musea v Brno 51: 261–270. 265–271. Young, D.A. & M. Beier., 1963a. Types of Cicadellinae (Homoptera: Cicadellidae) in the Natural History Museum in Vienna. – Annalen des naturhistorischen Received: 30 July 2007 Museums in Wien 67: 565–575. Accepted: 1 September 2007

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