Norwegian Journal of Entomology

Volume 47 No. 1 • 2000

Published by the Norwegian Entomological Society Oslo and Stavanger NORWEGIAN JOURNAL OF ENTOMOLOGY

A continuation of Fauna Norvegica Serie B (1979-1998), Norwegian Journal ofEntomology (1975­ 1978) and Norsk Entomologisk TIdsskrift (1921-1974). Published by The Norwegian Entomological Society (Norsk entomologisk forening). Norwegian Journal of Entomology publishes original papers and reviews on taxonomy, faunistics, zoogeography, general and applied ecology of insects and related terrestrial arthropods. Short com­ munications, e.g. less than two printed pages, are also considered. Manuscripts should be sent to the editor.

Editor Lauritz S~mme, Department of Biology, University of Oslo, P.O.Box 1050 Blindem, N-03l6 Oslo, Norway. E-mail: [email protected].

Editorial secretary Lars Ove Hansen, Zoological Museum, University of Oslo, Sarsgate 1, N-0562 Oslo. E-mail: [email protected].

Editorial board Ame C. Nilssen, Troms~ John O. Solem, Trondheim Uta Greve Jensen, Bergen Knut Rognes, Stavanger Ame Fjellberg, Tj~me

The goal of The Norwegian Entomological Society is to encourage the study of entomology in Norway and to provide a meeting place for those who are interested in the field. Annual membership fees are NOK 200 Guniors NOK 100) for members with addresses in Norway, and NOK 220 (Juniors NOK 110) for members abroad. Inquiries about membership should be sent to the secretary: Jan A. Stenl~kk, P.O.Box 386, N-4oo2 Stavanger. Norway. E-mail: [email protected].

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Subscription. Members of The Norwegian Entomological Society receive Norwegian Journal of Entomology and lnsekt-Nytt free. Institutional and non-member subscription NOK 250. Subscription and membership fees should be sent to the treasurer: Bj~m 0kland, Brekkeskog 31, N-1430 As, Norway; account no.: 08065440920.

Front cover: Boreophilus henningianus Sahlberg, 1832 (Coleoptera, Staphylinidae). Artist: Frode 0degaard.

Printed by: Reprografisk Industri AS, Oslo.

ISSN 1501-8415. Norw. J. Entomol. 47, 1-6. 2000

Cereals as host plants for thrips (Thysanoptera) in Norway

Sverre Kobro, Ann Elin Teksdal & Arild Andersen

Kobro, s., Teksdal, A. E. & Andersen, A. 2000. Cereals as host plants for thrips (Thysanoptera) in Norway. Norw. 1. EntomoI. 47, 1-6.

A survey of thrips (Thysanoptera) in cereals was carried out in southeastern Norway in 1996. No high densities of thrips were recorded. Limothrips denticornis and Frankliniella tenuicornis were the dominating species. Stenothrips graminum, which is new to the Norwegian fauna, was also found in abundance. 1. denticornis reproduced in the cereals only to a small extent. F. tenuicornis completed two generations, one in winter cereals in June, and a second generation in spring cereals in July. Pollen as an oviposition stimulant is discussed.

Key words: Thysanoptera, Limothrips denticornis, Frankliniella tenuicornis, Stenothrips graminum.

Sverre Kobro, Plant Protection Centre, Department ofEntomology and Nematology, Hegskoleveien 7, N-1432 As, Norway. Ann Elin Teksdal, Mdkevn. 2, N-4623 Kristiansand S, Norway. ArildAndersen, Plant Protection Centre, Department ofEntomology andNematology, Hegskoleveien 7, N-1432 As, Norway

INTRODUCTION tenuicornis, but H aculeatus may also cause se­ vere damage. TOOps (Thysanoptera) are often neglected as pests on outdoor cultivated plants in Norway, but dama­ 1. cerealium andH aculeatus were previously re­ ge has occasionally been ascribed to thrips. Crops ported as the most common pests in cereals in may be attacked by different species of tOOps in southern and central Sweden (Johansen 1946). different parts of the world (Lewis 1973). This Later Larsson (1995) stated that 1. denticornis and seems especially true for cereal tOOps in Europe. Stenothrips graminum Uzel, 1895 were also pests Haplothrips tritici (Kurdjumov, 1912) is dominant in Sweden, while populations of Thrips angusti­ on wheat in eastern and southern Europe. In wes­ ceps Uzel, 1895, A. obscurus and F. tenuicornis tern Europe the niche is filled by Limothrips cere­did not occur at injurious levels. aleum Haliday, 1836. Further north, in Scandina­ In Denmark 1. cerealium, 1. denticornis and H via, Limothrips denticornis Haliday, 1836, Ana­ aculeatus are the most important tOOps pests in phothrips obscurus (Muller, 1776), Frankliniella cereals (Anon. 1998), while F. tenuicornis is com­ tenuicornis (Uzel, 1895) and Haplothrips acule­ monly found, but rarely in damaging numbers atus Fabricius, 1803 are dominant in wheat (Koppa (Maltbrek 1932). Thus, 1. denticornis is the only 1967). F. tenuicornis and 1. denticornis have oats species regarded as a pest in all the three coun­ as their host plant in Britain (Pitkin 1976), but are tries, Denmark, Finland and Sweden, but 1. cere­ a minor problem in oats in Sweden (Larsson 1995). aleum is the most studied and described species Even within Scandinavia there are differences in (Johansen 1938). the choice of cereals as host plants for thrips. The knowledge on occurrence of tOOps living on Koppa (1970) focused on two species oftOOps as cereals and grasses in Norway is limited. Herstad pests in cereals in Finland, 1. denticornis and F. (1960) published some notes based mainly on in­ Kobro, Teksdal & Andersen: Cereals as host plants for thrips (Thysanoptera). fonnation from the government entomologists importance as pests on cereals in Norway, and WM. Sch0yen and T.H. Sch0yen for the years thereby contribute to later studies on damage 1862 to 1939. Some field trials have been perfor­ thresholds. med in cereals for control ofthrips in general (An­ dersen 2000), but there is little infonnation on identified thrips species as pests in cereals and MATERIALS AND METHODS grasses. In an experiment on chemical control, Chi­ Plant material was sampled in As, Akershus in rothrips manicatus Haliday, 1836, F. tenuicornis, southeastern Norway during the summer of 1996. L. denticornis and A. obscurus were found, but in One sample was taken once a week from each of small numbers (SeInes 1987). L. cerealeum is rare 14 cereal fields, where no insecticides had been and H aculeatus and T. angusticeps are not re­ used. The fields were in an open and flat agricul­ corded in Norway (Olsen & Solem 1982). tural area. The soil has a high content of clay and As the infonnation on occurence of thrips living the climate is semi-coastal. The cereal fields inves­ on grasses and cereals in Norway is limited, fur­ tigated were winter rye (n=2), winter wheat (n=2), ther infonnation must be obtained from our neigh­ triticale (n=2), barley (n=3), spring wheat (n=2) bouring countries, Denmark, Finland and Swe­ and oats (n=3). The results are given as averages den, where this insect group has been better stud­ of these and no further statistics are used. A redu­ ied for many years. ced number of samples (in rye and oats and every second week only) were taken in addition in 1998. The host plant choice of cereal thrips in Norway is also poorly known. Thrips have not evolved Sweep-netting has been the most common collec­ along phyletic lines ofhost plants in the same way ting method used, and the species composition as many other insect groups. The ancestor ofthrips presented in some papers (eg. Johansen 1938, was probably a detritus feeder, which later radi­ Koppii 1967) therefore contains several thrips spe­ ated onto the leaves and flowers of higher plants cies living in dicotyledons only. In this work we (Mound & Teulon 1995). There they have a var­ washed the collected plant material, a method ied diet, including pollen, on which they are de­ which is highly effective (Lewis 1973), thus avoi­ pendent for reproduction (Kirk 1995). A cereal leaf ding flowerthrips species living on weeds. can trap pollen from any plant in the vicinity, even The washing method is therefore better than from trees in a nearby forest, and thus provide food sweep-netting, but suboptimal for recording thrips and act as a potentional host plant for thrips. Such in cereals. Some species, L. denticornis in parti­ opportunistic behaviour is found for many thrips cular, spend most of their time within the leaf species (Mound & Teflon 1995), and may lead to sheaths, and may be trapped there during the wash­ different possibilities of host plant choice. ing process. In a method study, 60% of the ima­ The impact ofthrips as pests in cereals is difficult goes were recorded after washing, compared to to evaluate because of their vagile and opportu­ freezing the plant material. nistic habit, and reproductive demands. They may Each sample consisted of 25 straws cut at their land on a plant that they do not eat, or feed on it bases. The plant material was stored in plastic bags but not reproduce there (Mound 1997). Additio­ during transport to the laboratory, where water nally, too many other unknown factors influence with a drop of detergent was added. After at least whether or not an attack leads to damage (Skinner 30 minutes and gently shaking a few times, the & Parker 1995). plant material was removed from the bag, and the The current infonnation on occurrence and host water filtered through a nylon cloth (Kobro 1996). plant choice of the thrips species that can cause Thrips were stored in AGA (60% ethanol + glyc­ damage in cereals in Norway thus seems unclear. erol + acetic acid = 10 + 1 + 1) until identifica­ The aim of this study was to promote a better un­ tion. The numbers ofthrips were used as an indi­ derstanding of the occurrence of thrips and their cation of occurrence.

2 Norw. J. Entomol. 47, 1-6. 2000

Correct identification ofthrips may be complicated in the additional samples in rye and oats in 1998 due to an elaborate preparation technique and the (not shown) confirmed these results, except that need for a good microscope. The cereal thrips spe­ there were L. denticornis males in rye in June in cies recorded in Norway are very different mor­ 1998. phologically. As the collecting in our study was performed by the selective method of sampling and washing plant material, the thrips species can DISCUSSION easily be distingiushed by characters clearly vis­ No high number ofthrips was found as compared ible without preparation, in a binocular microscope to the threshold of damage suggested by Larsson with 40 times magnification. A simplified identi­ (1995). fication key has been made (Kobro 1997). Ima­ goes were identified according to this key, and the Imagoes ofL. denticornis appeared in similar num­ nymphs identified according to Koppii (1970) and bers in the winter cereals rye and triticale (Figure Larsson (pers. comm.). 1), but no nymphs were found. The abundance of imagoes indicates that two generations were com­ pleted, which was expected (Koppii 1970), but as RESULTS nymphs were found mainly in July and August in barley (Figure 2), most of the reproduction must Thrips were recorded in low numbers only. L. have occurred outside the cereal fields. However, denticornis andF tenuicornis accounted for 92.6% a low number of males were found in rye and ofthe total number of imagoes, and S. graminum triticale, and as the males are wingless, they must for 6% ofthe imagoes. Almost all nymphs found have developed on the sampled plants (Koppii belonged to these three species. The few pupae 1970), indicating that some reproduction must that were collected were not identified. Imagoes have taken place there. ofA. obscurus, Chirothrips hamatus Trybom, 1895 and C. manicatus were also recorded, but in low In the additonal samples of rye in 1998, however, numbers. a few nymphs and some males of L. denticornis were found in June. Thus there was reproduction L. denticornis. Occurrence ofimagoes was simi­ in rye in 1998, in contrast to 1996, and the repro­ lar in rye and triticale (Figure 1), while very low ductive behaviour of L. denticornis was different numbers were found in winter wheat (not shown). in the two years. This behaviour may have several No nymphs were found in winter cereals, while explanations, but might be a response to the pre­ barley contained some imagoes and a few nymphs sence or absence ofan oviposition stimulant (Terry in July and August (Figure 2). 1997) such as pollen. If they do not find the right R tenuicornis. A small peak of imagoes was re­ stimulus on a plant, thrips do not remain there. corded in May in the winter cereals (Figure 3), Such a selective behaviour must be important in while in July they were evenly distributed in win­ predicting consequences of an observed attack. ter cereals (Figure 3) and barley (Figure 4). The In barley, imagoes of L. denticornis also showed highest number ofnymphs were recorded in win­ a two-peaked occurrence possibly indicating two ter cereals in June (Figure 3) and in barley in July generations, but nymphs were found mainly in July and August (Figure 4). Only a few nymphs were (Figure 2). found in spring wheat and oats (not shown). For many thrips species the true host plant, that is S. graminum. In spite ofvery low numbers ofima­ where the breeding occurs, is not fully known. goes, this species represented the highest number Most records of «host plants» are only finding­ of nymphs in the cereals investigated and were places (Mound 1997). Our results indicate that rye found in oats (Figure 5). More than 95% of these and triticale had little importance as reproductive were S. graminum nymphs. hosts for L. denticornis. This contradicts the infor­ The occurence ofL. denticornis and S. graminum mation from Finland (Koppii 1970).

3 Kobro, Teksdal & Andersen: Cereals as host plants for thrips (Thysanoptera).

1.2

1.0

0.8

0.6

0.4

0.2 J r. I 11 1I I Figure 1. Imagoes of • Limothrips denticornis in rye and triticale.

1.2

1.0

0.8

0.6 ~ ca ... 0.4 en -... Figure 2. Imagoes and Q) 0.2 Q. nymphs of Limothrips ... denticornis in barley. Q) ..c E 1.5 :::J C C ca Q) 1.0 ~

0.5 Figure 3. Imagoes and nymphs of Frankliniella tenuicornis in winter cereals. 2.5 4~~~~------~

2.0

1.5

1.0

0.5 Figure 4. Imagoes and nymphs of Frankliniella tenuicornis in barley. 14 4 12 19 26 3 10 17 24 31 6 15 27 May June July August

4 Norw. J. Entomol. 47, 1-6. 2000

;: co.... 3.0 ­ rJ) -.... Dlmagoes Q) 2.5 ­ l a. .... I• Nymphs I Q) 2.0 ­ .0 E 1.5­ ~ c c LO- CO Figure 5. Imagoes and Q) :E 0.5 ­ nymphs of Stenothrips n n r r graminum in oats. 14 4 12 19 26 3 10 17 24 31 6 15 27 May June July August

Imagoes ofF tenuicornis had an early appearance needed on the distribution of the species and on ofoverwintered females, though in very low num­ oviposition preferance. bers in barley (Figure 4). Similar numbers ofima­ goes were seen in winter cereals and barley during Acknowledgements. We are grateful to Ricardo Hol­ July and August (Figure 3 and 4). In contrast to L. gado for inspiring discussions and for critical remarks denticornis, F tenuicornis reproduced twice in the on the manuscript. cereal fields, with a first generation in winter cere­ als (Figure 3) and a second one in spring cereals, References mainly in barley (Figure 4). This is well in agree­ ment with information from Finland (Koppa 1970) Andersen, A. 2000. Spr0)'ling mot trips i bygg. Gmnn and Sweden (Johansen 1938). forskning 1, 132-133. Anonymous 1998. http://www.planteinfo.dk/Informa­ Johansen (1938) stated that S. graminum is com­ tion/PCPlantevaern/skadedyr.html#2270. mon in Denmark, but rare in southern Sweden. Herstad, B. 1960. Opplysninger om forekomst av trips The species is presently common in southern Swe­ (Thysanoptera) i Norge. Norsk Ent. Tidsskr. 11, 145­ den (Larsson 1995), and is spreading northwards 149. (Larsson pers. comm.). It was first registered in Johansen, E. 1938. Studier mrande de pa gras och siides­ Norway in 1995 (Kobro unpublished), and is re­ slag levande tripsamas biologi och skadegorelse. ported here for the first time, and in considerable Statens Viixtskyddsanstalt, Meddelande Nr. 24, 1­ numbers (Figure 5). 65. Johansen, E. 1946. Studier och fOrsok rorande de pa griis och siidesslag levande tripsamas biologi och skade­ gorelse. (English summary). Statens Viixtskydds­ CONCLUSION anstalt. Meddelande Nr. 46, 1-59 The recorded numbers of tOOps were very low, Kirk, W. D. 1. 1995. Feeding behaviour and nutritional and far below a risk of damage in the cereals. requirements. Pp. 21-30 in Parker, B., Skinner, M. & Lewis, T. (eds.). Thrips biology and management. Reproduction ofL. denticornis was different from Plenum Press, New York and London. F tenuicornis and S. graminum, as all three spe­ Kobro, S. 1996. Notes on thrips (Thysanoptera) in Ice­ cies were abundant in cereals, but L. denticornis land. Fauna norv. Ser. B 43, 95-97. had a low reproduction rate there, in contrast to Kobro, S. 1997. Trips pa korn og gras. Norsk Landbruk the two other species. 11,13-15. Koppii, P. 1967. The composition of the thrips species To evaluate the true danger of injury by tOOps in in cereals in Finland. Ann. Agric. Fenn. 6, 30-45. cereals in Norway, more detailed information is Koppii, P. 1970. Studies on the thrips (Thysanoptera) species most commonly occurring on cereals in Fin­

5 Kabra, Teksdal & Andersen: Cereals as hast plants far thrips (Thysanaptera).

land. Ann. Agric. Fenn. 9, 191-265. Larsson, H. 1995. Trips i strasad. Faktablad om vaxt­ skydd, jordbruk 73. 4 pp. Sveriges lantbruksuni­ versitet, Uppsala. Lewis, T. 1973. Thrips, their biology, ecology and eco­ nomic importance. 349 pp Academic Press, London and New York Maltb

Received 19 August 1999. accepted 15 November 1999

6 Norw. J. Entomof. 41, 7-19. 2000

Contribution to the knowledge of Norwegian Coleoptera

Frode 0degaard & Sindre Ligaard

0degaard, F. & Ligaard, S. 2000. Contribution to the knowledge ofNorwegian Co1eoptera. Norw. J. Entomol. 47, 7-19.

The following 34 species of Co1eoptera are reported as new to Norway: Hydroporus nigellus Mannerheim, 1853, Bembidionfumigatum (Duftschmid, 1812), Dromius linearis (Olivier, 1795), Leiodes inordinata (ISah1berg, 1898), Colon pseudolatum Palm, 1941, Omalium littorale Kraatz, 1858, Bibloplectus tenebrosus (Reitter, 1880), Bibloplectus spinosus Raffray, 1914, Stenus latifrons Erichson, 1839, Stenus ochropus Kiesenwetter, 1858, Philonthus pseudoparcus Brunne, 1976, Mycetoporus eppelsheimianus Fage1, 1965 (brucki auct. nec Pandelle, 1869), Philhygra pinegensis Muona, 1983, Falagrioma thoracica (Stephens, 1832), Gyrophaena transversalis Strand, 1939, Placusa suecica Johnson & Lundberg, 1977, Cypha suecica (Palm, 1936), Ernobius pini (Stunn, 1837), Meligethes tristis Stunn, 1845, Psammoecus bipunctatus (Fabricius, 1792), Enicmus lundbladi Palm, 1956, Stilbus atomarius (Linnaeus, 1767), Stilbus oblongus (Erichson, 1845), Meloe proscarabaeus Linnaeus, 1758, Bruchus affinis Frolich, 1799, Longitarsus curtus (Allard, 1860), Asiorestia transversa (Marsham, 1802), Asiorestia interpunctata (Motschu1sky, 1859), Apion columbinum Gennar, 1817, Apion melancholicum Wencker, 1864, Barynotus moerens (Fabricius, 1792), Coeliodes nigritarsis Hartmann, 1895, Gymnetron labile (Herbst, 1795), Taphrorychus bicolor (Herbst, 1793). Notes on ecology and distribution in the neighboring countries are given for each of the species. Ampedus rufipennis (Stephens, 1830) is deleted as a Norwegian species. Some possible explanations for the continuous addition of new species to the Norwegian fauna are discussed.

Key words: biogeography, Co1eoptera, geographical distribution, faunistics, new species in Norway.

Frode 0degaard, Norwegian Institute for Nature Research, Tungasletta 2, N-7485 Trondheim, Norway. Sindre Ligaard, Mads vei 21, N-1540 Vestby. Norway.

INTRODUCTION circumstances around the records in such publi­ cations are often hardly available or incomplete. The beetle fauna ofNorway is far from completely Thus, we find it important to publish separate notes known. The number of beetle species recorded on species new to the country. The material is from Norway has increased from 3091 species in mostly collected during the last decade. Unless 1961 to 3430 species in 1997 (Bakke 1961, Hans­ anybody else is mentioned, the identifications of sen et al. 1997). Accordingly, more than 9 new the material are performed by the authors. The no­ species are registered every year on average (339 menclature follows Lawrence & Newton (1995) species/36 years). The estimated number of Nor­ at family level and Lundberg (1995) at species wegian beetle species is 3800 (Ottesen 1993). This level. paper summarizes data on 34 beetle species pre­ viously not published as new to Norway although The following abbreviations are used in the text: some of the species are mentioned in reports or BT=Barber trap, WT=Window trap, F0D= Frode other sources (e.g. Vik 1995, Hanssen et al. 1997, 0degaard, NINA=Norwegian Institute for Nature T0mmeras et al. 1997, Hanssen & Hansen 1998, Research, OHA= Oddvar Hanssen, SLI=Sindre Direktoratet for Naturforvaltning 1999). Data and Ligaard, 2MB=Zoologica1 Museum, University of

7 /Jdegaard & Ligaard: Contribution to Norwegian Co/eoptera

Bergen, ZMO=Zoological Museum, University of Hl:Il1sen 1996). If there are breading populations Oslo. Provincial abbreviations follow 0kland in Norway, the species has to be considered as vul­ (1981). nerable (Direktoratet for Naturforvaltning 1999).

Dromius /inearis (Olivier, 1795) DYTISCIDAE We can report new records of Dromius linearis from several Norwegian localities: VAY Lyngdal: Hydroporus nige//us Mannerheim, 1853 Korshavn (EIS 2), 23 July 1992, leg. & colI. SLI o Hvaler: Asmal0y, Viker (EIS 12), several exx., & F0D. Two specimens found among dense leg. & colI. SLI. Hvaler: 0rekroken (EIS 12), one vegetation in a salt marsh dominated by Triglochin ex., leg. & colI. SLI. Fredrikstad: Torsnes (EIS maritima, Plantago maritima and grasses (Poa­ 20), several exx., leg. & colI. SLI. All specimens ceae). This habitat is not typical for D. linearis were collected in coastal rock pools. whose normally occurs in dunes at coast (Lindroth 1986). However, the species apparently reproduce The H. nigellus-complex was revised by Nilsson at the locality because ten individuals were found (1994). H. nigellus is hard to separate from H. during a revisit in May 1994, leg & colI. SLI. 0 geniculatus, however, their ecology is very diffe­ Hvaler: 0rekroken (EIS 12), 28 May 1993, leg. rent. While H. geniculatus is a common species F0D. Two specimens found among scattered vege­ in temporary pools in higher altitudes (frequently tation in sand dunes. In addition, one specimen occurring above the tree line), H. nigellus is restric­ was observed at the same locality 9 July 1997 (l ted to coastal rock pools with fresh water (Nilsson Andersen pers. comm.). VE Tj0me: Moutmarka & Holmen 1995). The species is expected to be (EIS 19), 7 Sept. 1993, leg. & colI. OHA. TEY more widely distributed in Norway due to the Kragem: Straholmen (EIS 11), June 1993, leg. & recent clarification ofthe species complex. colI. OHA. At both localities, several specimens were found by beating stems ofRumex crispus on CARABIDAE gravelly ground (OHA pers. comm.). One old, record of D. linearis is reported from AK Oslo, Bembidion fumigatum (Duftschmid, 1812) T0yen, leg. N. G. Moe (Sch0yen 1879). However, VE Tj0me: Moutmarka (EIS 19), 29 May 1993, this record was considered as uncertain by Th. leg. & colI. F0. One single female was found in a Munster and A. Strand (Hellen 1939). Regarding salt marsh at the edge of a pool with rich vegeta­ the collection of N.G. Moe, see Miinster (1923­ tion. 24, pp. 267-268). It is unknown whether the record represents a per­ The localities for this species in the Oslofjord-area manent population or an accidentally introduced are of very high conservation value (Hanssen & specimen. The specimen was found in its natural Hansen 1998). D. linearis is common and wide­ habitat, a fact that indicates a wider occurrence of spread in Denmark and south Sweden and, thus, the species. However, several collectors have sear­ the species was expected to occur in Norway. ched for the species at the same locality without However, it is somewhat remarkable that the spe­ luck. On the other hand, the species is now and cies was found at four separate localities during a then collected in wind drift at seashores (Lindroth period of two years. Especially, because some of 1985). One hypothesis is that the Tj0me-specimen the localities were very well investigated during has arrived with wind drift and eventually found the 1980s without any record of this easy trace­ its natural habitat in the vicinity. This record is able species. Therefore, it seems reasonable to mentioned in Hanssen & Hansen (1998) as a repre­ propose that the species has expanded its range. If sentative of a locality of very high conservation the old report from T0yen is true, it may indicate value. The species is common and widely distri­ a wider range at that time. At a larger temporal buted in Denmark, but it is only known from the scale, the range of the species may fluctuate furthest south in Sweden (Sk, Bl) (Lundberg 1995, continuously in accordance with climatic changes.

8 Norw. J. Entomol. 41, 1-19. 2000

The present localities should be revisited in order traps were situated in a salt marsh. () Hvaler: to see if the species has established pennanent 0rekroken (EIS 12),6 June 1995, leg. & colI. SLI. populations, or if the records only represent tem­ Several individuals collected under wrack beds poral fluctuations in the outer margins ofits range. (sea-weeds) on sandy banks. 0. littorale was earlier considered as a variety of 0. caesum (Palm 1948), and the status ofthe spe­ LEIODIDAE cies was debated in the 1950s (Palm 1962). Leiodes inordinata (J.Sahlberg, 1898) Eventually, Lohse (1960) showed that 0. littorale is a valid species. For identification, see Palm NTI Lierne: Skograudberga (EIS 103), WT 17 July (1962), Freude et al. (1964) or Palm & Lundberg - 21 Aug. 1986,1 ex., leg. & colI. OHA. The trap (1993). The species is restricted to wrack beds was situated in a south faced gap in an old spruce (Palm 1948) and thus, it is more specialised than forest. Mosvik: Kilen (EIS 97), BT 1-24 July 1994, its closest relatives. There are scattered records of 1 ex., leg. NINA, colI. F0D. The trap was situated the species along the Swedish coastline (Lundberg in an old spruce forest (T0mmeriis & Breistein 1995). 1995, T0mmeriis et al. 1997). This is one ofthe most characteristic Leiodes-spe­ Bibloplectus tenebrosus (Reitter, 1880) cies due to the irregular rows of puncture of the AK Nes: Israelsmosen (EIS 37), BT Aug. 1992, 1 elytra (Strand 1957). In Sweden, the species is ex., leg. & colI. F0D. The trap was situated in an distributed in the boreal zone from Vannland and ombrotrophic mire. northwards (Lundberg 1995), where the species In Sweden, there are rather few localities of this is nonnally caught in traps in dense forests (Stig species. It dominates in the southern provinces, Lundberg pers. comm.). Due to the subterranean however, it is recorded as far north as Norrbotten way ofliving, the Leiodes-species are rarely collec­ (Lundberg 1995). There is a certain probability ted and most species are probably strongly over­ that the species have been overlooked in Norway looked. These relations and the fact that the ground due to the small size and the rather concealed habit. fauna ofdense forests, such as bilberry woodland, Moreover, mire and fen habitats are not satisfac­ are poorly investigated, makes it reasonable to beli­ tory investigated regarding invertebrates. eve that L. inordinata is continuously distributed throughout the Southern and Middle boreal zone Bibloplectus spinosus Raffray, 1914 in Norway. () Hvaler: Arekilen (EIS 12), 28 May 1993, leg. Colon pseudolatum Palm, 1941 & colI. F0 & SLI. Two specimens were collected in wet mosses at the shore of an eutrophic pool FI Karasjokk: Amujiikka (EIS 159), BT July -Aug. dominated by Phragmites-vegetation. 1996, 5 exx., leg. H. Rinden, colI. SLI. This record is mentioned in Hanssen & Hansen The species belongs to the Northern boreal vege­ (1998) as a representative ofa locality ofvery high tation zone, where it is found among grass roots conservation value and the species is considered on dry, gravelly ground (Palm 1941). The species as declining and care demanding in Norway (Di­ is known from northern Sweden southwards to rektoratet for Naturforvaltning 1999). The species Halsingland and from Finland (Lundberg 1995). was expected to occur in Norway because it is widely distributed in southern parts of Sweden, and it is also known from the adjacent province STAPHYLlNIDAE Vannland (Lundberg 1995). Omalium littorale Kraatz, 1858 Stenus latifrons Erichson, 1839 VE Tj0me: Hvasser, S0nstegiird (EIS 19), BT Aug. () Hvaler: Arekilen (EIS 12), 19 May 1990,1 ex., 1985, several exx., leg. T. Andersen, colI. SLI. The leg. & colI. SU, 28 May 1993,4 exx., leg. & colI.

9 ft:}degaard & Ligaard: Contribution to Norwegian Coleoptera

F0, 28 June 1993, several exx., leg. & colI. SLI. 1992, Lundberg 1995). Accordingly, this species All specimens collected at the shore of an eutro­ needs to be added to the Norwegian list ofColeo­ phic pool dominated by Phragmites-vegetation. ptera. This is the first verified record ofthis species from Mycetoporus eppe/sheimianus Fagel, 1965 NOIway. There is a report ofS. latifrons from NSI, (brucki auet. nee Pandelle, 1869) StOIjord (leg. Hageman) (Strand 1946). However, MRI Sunndal: Gravem (EIS 79), WT 2 June - 12 this specimen could not be found in any of the July 1986, 1 ex., leg. & colI. OHA. The trap was Norwegian museums. The report from NSI is pro­ situated in a deciduous forest dominated by aspen bably a mistake and should therefore be omitted (Populus tremula) in a steep, south-faced slope. (Torstein Kvamme pers. comm.). The records from ON Nord-Fron: Hesteskobakken (EIS 62), 11 July Arekilen are mentioned in Hanssen & Hansen 1988, leg. & colI. F0D, det. J. Muona. One speci­ (1998) as representatives ofa locality ofvery high men sifted from decaying leaves in an old alder conservation value. The species is considered as forest. AK Asker: Skaugurnasen (EIS 28), WT July declining and care demanding in Norway (Direk­ 1991, 1 ex., leg. J. Stokland, colI. F0D, det. J. toratet for Naturforvaltning 1999). Muona. The trap was situated in a thermophilous, Stenus ochropus Kiesenwetter, 1858 deciduous forest. TEI Seljord: Heggeneset (EIS 17), BT 8 April ­ The species is normally found among decaying 30 May 1998, 1 cJ and 1 9, BT 25 July - 6 Sept leaves and mosses on the forest-floor (Palm 1966). 1998, 1 9, leg. & colI. F0D. The traps were put In Germany, the species is found among decaying out in steep and open slopes towards the lake, leaves at the basis of old oaks, in rotten stumps, Seljordsvannet. and especially in Polyporus-fungi (Koch 1989a). It is scattered distributed in southern parts ofScan­ This record is somewhat surprising because the dinavia and evidently rare everywhere (Lundberg nearest localities are situated in SE Sweden, where 1995, Hansen 1996). The species has to be consi­ the species is found on dry, sandy or gravelly dered as declining and care demanding in Norway calcareous soils, often on so called «Alvarmark» (Direktoratet for Naturforvaltning 1999). (Palm 1961). The hilly topography in Telemark county, causing extremely warm microclimate in Phi/hygra pinegensis Muona, 1983 the south faced slopes at summertime. In this lo­ F0 Sor-Varanger: Nyrudmyra (EIS 160), 8 July cality natural stone- and soil-slides keep the loca­ 1998, leg. & colI. F0. One female sifted from drift lity open and sun exposed. Combined with calcare­ material washed up after a flood in a small river. ous soil types, such localities are probably functio­ The vegetation in the flooded area was dominated nally very similar to the areas in SE Sweden. The by willow thickets (Salix spp.), Betula nana, record ofS. ochropus is probably another example Carex-spp and grasses (Poaceae). The locality lies ofrelicts from warmer periods in post-glacial time in association with a larger bog-complex in the in Norway. The species has to be considered as 0vre-Pasvik area in Finnmark. declining and care demanding in Norway (Direk­ toratet for Naturforvaltning 1999). This species was described as late as in 1983 after one male from Pinega in NW Russia (Muona Phi/onthus pseudoparcus Brunne, 1976 1983). Later, several individuals were found in The describtion ofPhilonthuspseudoparcus is ba­ Oulanka National Park in Kuusamo, Finland sed, among others, on 35 specimens from AK (Muona 1995). The female is described by Muona Asker: Bmnnoya (EIS 28), June 1967, and June (1995). Due to many independent records the last 1971, leg. A. Strand (Brunne 1976). However, the 20 years, there is reason to believe that the species species has never been registered as Norwegian has expanded its range towards the west recently. in the catalogues on Nordic Coleoptera (Silfver­ On the other hand, range expansions are a rare berg 1979, Lundberg 1986, Vik 1991, Silfverberg phenomenon in natural habitats. Accordingly, it

10 Norw. J. Enlomo/. 41, 1-19. 2000 will be interesting to see further reports on the spe­ Niederosterreich (Strand 1939). Later, it is only cies. recorded from Hamburg, Rheinland in Germany, and from The Czech Republic. The species can Falagrioma thoracica (Stephens, 1832) easily be identified in Strand (1939). In Central TEI Seljord: Heggeneset (EIS 17), BT 30 May ­ Europe, the species is collected in the fungi Boletus 6 Sept. 1998, ca. 100 exx., leg. & colI. F0D. scaber (Koch 1989a). The Norwegian specimens Although the species was abundant throughout the were found under decaying leaves on silty ground summer, no specimens were caught in the period in the spring, a habitat that probably represents from 8 April to 30 May, indicating that the species the hibernating site ofthe population. The species is a summer breeder. is mentioned in Andersen & Hanssen (1994) as a representative ofthe rich and peculiar insect fauna The species is only recorded from Skane and Got­ along the riversides of Gaula. The species has to land in Sweden (Lundberg 1995), although in Den­ be considered as vulnerable in Norway (Direkto­ mark it is recorded from most provinces (Hansen ratet for Naturforvaltning 1999). 1996). Similar to the present record, Palm (1968) reports that the species often occurs in large num­ Placusa suecica Johnson & Lundberg, 1977 bers. Elsewhere in Scandinavia, the species is NTI Mosvik: Kilen (EIS 97), WT June - Sept. 1994 mostly collected near the coast under sea-weeds and 1996, 4 exx., leg. NINA, colI. F0D. The traps or in plant roots on moist, sandy soils (Hansen were situated in an old coniferous forest dominated 1954, Palm 1968). In this regard, the Telemark­ by Norway spruce (Picea abies) (Tl'lmmeras & record is very strange. Obviously, its occurrence Breistein 1995, Tl'lmmeras et al. 1997). in Norway has to be seen in association with the relict argument proposed for Stenus ochropus (see In Sweden, the species is collected in window traps above). In addition, there are probably unrevealed around timber-piles (Johnson & Lundberg 1977), facts attended with the ecology of the species and at present, it is recorded from several provinces which tie together the different circumstances of in Central Sweden (Lundberg 1995). Placusa­ the records. Koch (1989a) reports that the species species are fungivorous in subcortical habitats is hygrophilous in northern parts ofits range while (Ashe 1991). The biology ofP. suecica is probably thermophilous elsewhere. The species is also found similar to that of congeneric species (Lundberg in association with ants (Koch 1989a). The species 1995). The species is apparently rather common has to be considered as declining and care deman­ and widespread in Scandinavia. Perhaps it was ding in Norway (Direktoratet for Naturforvaltning overlooked before window traps became common­ 1999). ly used for collecting swarming insects.

Gyrophaena transversalis Strand, 1939 Cypha suecica (Palm, 1936) STI Melhus: Melhus (EIS 92), 12 April 1992, leg. "Hvaler: Arekilen (EIS 12), 19 May 1990, leg & & colI. F0D. Several specimens were sifted from colI. SLI, and 28 May 1993, leg. & colI. SLI & old, dry leaves of Tussilago farfara lying on silty F0D. Several specimens collected among wet banks ofthe river Gaula. One specimen was caught mosses and Phragmites-vegetation at the shore of at the same locality by car-netting 18 May 1993, an eutrophic pool. leg. & colI. F0D. Several individuals collected by The distribution of this species in Scandinavia is sifting from decaying leaves of Salix triandra on restricted to central parts of Sweden (Palm 1966, silty ground at the same locality 21 April 1996, Lundberg 1995). Thus, the Norwegian occurrence leg. & colI. OHA. Furthermore, one specimen was ofthis species is apparently not surprising. How­ found in decaying drift material washed up at the ever, the species is also recorded in Denmark riverside of Gaula at STI Melhus: Udduvoll bru, recently (Hansen et al. 1991). In that regard, it is 10 June 1997, leg. & colI. F0D. likely that the species has expanded its range. The The species is new to the Nordic countries, and it record from Arekilen is mentioned in Hanssen & was described after 13 specimens collected in Hansen (1998) as a representative of a locality of

1 1 {}degaard & Ligaard: Contribution to Norwegian Co/eoptera very high conservation value. The species is con­ in the first half ofthis century, it is not reasonable sidered as declining and care demanding in Nor­ that the species has been overlooked by previous way (Direktoratet for Naturforvaltning 1999). collectors. Probably the species has expanded its range towards the north recently. In southern Swe­ den the species is common and a new northern ELATERIDAE limit was reported from Vastmanland recently (Erichson 1999). The climatic conditions in these Delete Ampedus fufipennis (Stephens, 1830) areas are not very different from the localities in The only report on this species from Norway is Norway, so perhaps the only hurdle for the pro­ from TEY Porsgrunn: Eidanger (EIS 11), leg. Th. gress ofthis species is the distribution of the host Munster, colI. ZMO. This specimen, however, is plant. Remarkably, Meligethes planiusculus (Heer, a misidentified Ampedus hjorti, det. F0D 1998. 1841) another congeneric species associated with A. rnfipennis should, therefore, be deleted as a Nor­ Echium vulgare, has expanded in Denmark and wegian species. Consequently, TEY should be Sweden recently (Hansen et al. 1992, Erichson added as a new provincial record for A. hjorti that 1999). M. tristis is considered as declining and also recently is recorded from the same province care demanding in Norway (Direlctoratet for Natur­ (Bakke 1999). forvaltning 1999).

PTINIIDAE SILVANIDAE

Ernobius pini (Sturm, 1837) Psammoecus bipunctatus (Fabricius, 1792) o Hvaler: 0rekroken (EIS 12),28 May 1993, leg. AK Oslo: 0stensjovann (EIS 28), 15 Aug 1992, & colI. F0D. Three specimens were beaten from leg. & colI. SLI. Several specimens collected Scots pine (Pinus silvestris). The pine-trees grew among decaying Phragmites-straws at the margin in scattered stands on open established sand-dunes. ofan eutrophic lake. AKVestby: Kjennstjern (EIS The next year several specimens were recorded in 28),20Aug. 1996, leg. & colI. SLI. Five specimens pine-cones at the same locality (leg. & colI. SLI). collected in swampy vegetation. 0 Aremark: This record is mentioned in Hanssen & Hansen Gjolsjo (EIS 21), 14 May 1994, leg. & colI. SLI (1998) as a representative ofa locality ofvery high & F0D. Several specimens collected among de­ conservation value. The Ernobius species develop caying Phragmites-straws at the margin ofa meso­ in cones or in thin twigs ofconiferous trees (Freude trophic lake. et al. 1969). A systematic review of the genus is The records from 0stensjovann and Gjolsjo are given by JoOOson (1966). In Sweden, the species mentioned in Hanssen & Hansen (1998) as repre­ is found northwards to Dalarna (Lundberg 1995). sentatives ofa locality ofhigh conservation value. The species is probably associated with decaying Phragmites where it is normally found (Hansen NITIDULIDAE 1964). In Sweden, the species is recorded north­ wards to Viirmland (Lundberg 1995). Me/igethes lristis Sturm, 1845 AK Brerum: Ostoya (EIS 28), 24 June 1996, leg. & colI. SLI. Brerum: Fornebu (EIS 28), 9 July 1997 and 9 June 1998, leg. & colI. F0D. Oslo: Gress­ CORTICARIIDAE holmen (EIS 28), 18 June 1999, leg. & colI. F0D. Enicmus /undb/adi Palm, 1956 Several exx. found at each locality. The species is reported from YE in Vik (1995). This species seems to be obligatory on its host The details around this record are the following: plant, Echium vulgare, in the inner parts of Oslo­ VE Tjome: Mostranda (EIS 19), 30 June 1985, fjorden. Since this area is rather good investigated

1 2 Norw. J. Enlomo/. 41, 1-19. 2000 leg. & coil. SLI. One specimen sweep-netted in The many recent records indicate that this species shrub-vegetation. is well established at least in the outer Oslofjord­ area. The species is also common and widespread All stages of this species are found in large amo­ in southern parts of Sweden. unts under the thin outer bark of recently broken trunks of aspen (Popu/us tremu/a) (Lundberg 1964). The species lives in association with the MELOIDAE fungi Hypoxy/on mammatum (Wg) Millar (Pyre­ nomycetes) which gives the bark a characteristic Meloe proscarabaeus Linnaeus, 1758 black appearance in the point of attack. This may SFI Lrerdal: Bjorkurn (EIS 51), 5 June 1995, leg. be seen from long distance, and it resembles inju­ & coli. SLI. One dead specimen collected under a ries from fire (Lundberg 1969). There are scattered stone in a south faced slope with scattered vegeta­ records ofthis species in southern parts ofSweden tion cover. and in Finland (Lundberg 1995). The species is considered as declining and care demanding in According to Andreas Strand's diary (ZMB), there Norway (Direktoratet for Naturforvaltning 1999). are uncertain records of M proscarabaeus from TEY Kragero, leg. E. Berg, 0 Halden, leg. Printz, AK Oslo, leg. Siebke. These records are probably PHALACRIDAE the basic data for the uncertain records from Nor­ way reported in Lindroth (1960). In ZMO, there Stilbus atomarius (Linnaeus, 1767) are three labeled specimens ofM proscarabaeus, o Riide: Visterflo (EIS 20), 29 Aug. 1992, leg. & det. SU, leg. Siebke, Esmark and Berg, respec­ coil. SLI. Five specimens collected in very wet tively. Specifications oflocalities are lacking, how­ vegetation dominated with Sparganium spp. at the ever. We expect that the specimens of Siebke and shore of an eutrophic pool. Berg are identical with those seen by A. Strand. The uncertainty attended with these records is, Little is known about the biology ofSti/bus-species therefore, probably due to the lack of locality but maybe they feed on algae (Koch 1989b). S. reports rather than to the identifications. In 2MB, atomarius is always found in association with .there are three additional specimens of M pro­ freshwater-shore vegetation. Either on grasses, scarabaeus, det. OHA, F0D, labeled B0 Modum, flowers, or in decaying plant material (Koch leg. N. G. Moe. These specimens are never 1989b). The occurrence ofS. atomarius in Norway reported from Norway, and it is hard to know whe­ is not surprising, because the species is widespread ther they are overlooked or disregarded by former in southern parts of Sweden. It is also recorded collectors. According to Miinster (1923-24, pp. from the Swedish provinces adjacent to SE Nor­ 267-268), several of the specimens collected by way. N.G. Moe are doubtful, due to insufficient labeling Stilbus oblongus (Erichson, 1845) and uncritical mixing of Norwegian and foreign material. Consequently, we choose to consider the VE Tonsberg: Prestrodkilen (EIS 19), 14 Oct. specimens from Modum as uncertain. All the 1984, 1 ex., leg. & coil. OHA. Tjome: Sunnane uncertain specimens are probably collected during (EIS 19), 10 July 1985, 1 ex., and 14. June 1987, the 19th century due to the main activity periods I ex., both leg. & coli. SLI. T011sberg: Akersvannet ofthese collectors. (EIS 19), 14 June 1989, I ex., leg. & coil. SLI. Stavern: Lydhusstranda (EIS 19),24 June 1989,4 The species is known to parasite the bee-species ex., leg. & coil. F0D. Tjome: Kulebekkilen (EIS Anthophora retusa, A. acervorum, and Andrena 19), 28 July 1992, 1 ex., leg. & coil. F0D. 0 jacobi (Hansen 1973). M. proscarabaeus is recor­ Hvaler: Arekilen (EIS 12),28 May 1993, I ex., ded from all Danish provinces (Hansen 1996), and leg. & coli. F0D. All specimens are found in northwards to Dalarna in Sweden. The species is association with Phragmites-vegetation. considered as vulnerable in Norway (Direktoratet for Naturforvaltning 1999). 1 3 edegaard & Ligaard: Contribution to Norwegian Co/eoptera

CHRYSOMELlDAE Asiorestia interpunctata (Motschulsky, 1859 ) Bruchus affinis Frblich, 1799 AK Fet: Nordre 0yeren (EIS 29),9 Aug. 1996, () Hvaler: 0rekroken (EIS 12),28 June 1993, leg. leg. & coil. F0D. One 9 sweep-netted on swampy SLI, coli. SLI & F0D. Five individuals recorded wetlands dominated by large Carex spp. and on Lathyros japonicus. AK Asker: Vollen (EIS grasses (Poaceae). 28), 26 July 1998, 1 ex., leg. & coil. F0D. One individual sweep-netted in calcareous meadow This species was published as new to Norway by vegetation. TEI Seljord: Heggeneset (EIS 12),8 Strand (1965), however later, the record was July 1997,30 May 1998, and 25 July 1998, leg. deleted for unknown reasons (Strand 1977). A & coil. F0D. revision ofthis specimen showed that it was an A. transversa (see above). A. interpunctata is not Collected in large amounts in reproductive parts recorded from Sweden (Lundberg 1995), but two of the host plant at the site, Lathyros silvestris, recent records are registered from Denmark (Han­ which was growing in large stands on base-rich sen et al. 1992, Hansen et al. 1993). The species soils along a roadside. The locality is south-faced is considered as declining, care demanding in Nor­ and very warm. B. affinis breeds in Lathyros spp. way (Direktoratet for Naturforvaltning 1999). and are probably spreading in Northern Europe (Hansen 1988). BRENTIDAE Longitarsus curtus (Allard, 1860) ON Nord-Fron: Hesteskobakken (EIS 62), 20 July Apion co/umbinum Germar, 1817 1994, I ex., BT 9 Sept. - 10 Oct. 1998, 2 exx. Sel: TEI Seljord: Heggeneset 8 July 1997, 30 May Otta, Solgjem (EIS 62), 21 July 1993, 7 exx., leg. 1998, 25 July 1998, 6 Sept. 1998, BT 8 April - 30 & coil. F0D. SFI Luster: Ornes (EIS 50), 28 Sept. May, and BT 30 May - 25 July 1998, leg. & coil. 1991, I ex., 29 May 1992, I ex., 15 Sept.l992, I F0D. The species was found in large amounts on ex., leg. & coil. F0D. its host plant, Lathyros silvestris, which occurred The species is oligophag on Boraginaceae, and it in large stands along a roadside on base-rich soils. is known from Symphytum spp., Pulmonaria spp. In the Nordic countries, A. columbinum is mono­ (Koch 1992), and Mysotis arvensis (Hansen 1964). phagous on the host plant and the larvae develop In Gudbrandsdalen (ON), we have found the spe­ in rolled leaflets (Gonget 1997). The species is cies on Lappula sp. in dry meadows on base rich very rare and local in Scandinavia, but often abun­ sandy soils. dant where it occurs. The species is known from a few localities in southern Sweden and one loca­ Asiorestia transversa (Marsham, 1802) lity in Denmark in recent times (Gonget 1997). () Hvaler (EIS 12), 27 June 1915, I 9, leg. The species has to be considered as declining and Hanssen, coil. 2MB, det. F0D. AK Brerum: care demanding in Norway (Direktoratet for Fornebu (EIS 28), 27 July 1998, leg. & coil. F0D. Naturforvaltning 1999). One single female sweep-netted on Cirsium arvense. Apion me/ancho/icum Wencker, 1864 The preferred host plant of A. transversa is C. TEI Seljord: Heggeneset, 8 July 1997, 30 May arvense, but the species is also found on C. 1998,6 Sept. 1998, leg. & coll. F0D. Found toget­ oleraceum and Carduus acanthoides (Hansen her withA. columbinum on the host plant, Lathyros 1929, Koch 1992). The species is recorded north silvestris. Although several specimens were found, to Dalarna in Sweden (Lundberg 1995). The A. melancholicum was not as numerous as A. Norwegian occurrence ofA. transversa was, there­ columbinum. fore, expected when the Swedish distribution was The species is monophagous on the host plant in extrapolated westwards. the Nordic countries, and the larvae develop in

1 4 Norw. J. Enlomo/. 47, 7-19. 2000 young fruits (G0nget 1997). The distribution an omprotrophic bog with dominance of Sphag­ pattern and occurrence ofthis species is very much num spp. in wetter parts. like that of A. columbinum. However, we have Whilst C. rubicundus is a generalist on Betula spp. seen no report on that the two species occur to­ in different habitats, C. nigritarsis is restricted to gether. A. melancholicum is only known from two bogs (Heijerman 1993). In Central Europe, the Danish (SZ, NWZ) and two Swedish (Sk, Up) species is monophagous on Betula pubescens provinces (G0nget 1997). The records ofthese two where the larvae probably develop in female cat­ Apion-species confIrm the potential ofthese areas kins (Koch 1992). The species is recorded north­ of Norway for inhabiting northern outpost popu­ wards to Tome Lappmark in Sweden (Lundberg lations of heat-loving species with distribution 1995). A more closely investigation of the bog centers much longer south. The species is consi­ systems in eastern parts of the country will pro­ dered as declining and care demanding in Norway bably reveal that the species is more widespread (Direktoratet for Naturforvaltning 1999). in Norway.

Gymnetron /abile (Herbst, 1795) CURCULlONIDAE {) Hvaler: Asmal"Y (EIS 12),21 July 1995, 1 ex., Barynotus moerens (Fabricius, 1792) leg. & col1. SU, primo Aug. 1995, 8 exx., leg. & col1. SU. The specimens were sweep-netted on HOY Bergen: Sverresborg, Skuteviken, Sand­ Plantago lanceolata in a salt marsh. viken (EIS 39), 19 and 22 June 1996, leg. J. Skartveit, col1. 2MB. Two individuals were found The species develops in stem galls or in the stem among dense vegetation ofshrubs and weeds (e.g. base of Plantago lanceolata (Hansen 1965). G. Rubus idaeus, Urtica dioica, Aegopodium poda­ labile is recorded in southern Sweden northwards graria, and Anthriscus sylvestris) in a small park to Bohusliin and Viistmanland (Lundberg 1995). surrounded by close settlements (l Skartveit pers. The species is considered as a declining, monitor comm.). {) Spydeberg: Hemnes (EIS 29), BT Aug. species in Norway (Direktoratet for Naturforvalt­ 1996, 1 ex., leg. & col1. SLI. The trap was situated ning 1999). in a small bog close to the lake Lyseren. Taphrorychus Nc%r (Herbst, 1793) The species is polyphagous on different herbs, and RI Suldal: Lomatjern (EIS 24), WT 24 May - 25 it is most commonly recorded in deciduous forests. June 1991, 1 ex., leg. 1. Stokland, col1. F0. The The adults are active at night and are normally trap was situated in a west-faced forest dominated found under stones or at the base of plants at with Betula pubescens, but also with occurrences daytime (Palm 1996). B. moerens is local and rare ofFraxinus exelsior, Sorbus aucuparia, andAlnus in Denmark and southern parts of Sweden (Palm incana. RI Suldal: Sandvika (EIS 24), WT 25 June 1996). The species is considered as a declining, - 25 July 1991, 1 ex., leg. J. Stokland, col1. F0. monitor species in Norway (Direktoratet for The trap was situated in woodland dominated by Naturforvaltning 1999). Scots pine (pinus silvestris). Betulapubescens and Coe/iodes nigritarsis Hartmann, 1895 Sorbus aucuparia occurred scattered between the Scots pines. The two localities lie 7 km from each STI Oppdal: Nerskogmyra (EIS 86), 13 June other (Stokland pers. comm.). 1993, leg. & col1. F0D. Two (J(J and one 9 were sweep-netted among Salix spp. Betula nana, and The primary host tree of T. bicolor is Fagus B. pubescens at the margins ofa rich fen. The loca­ silvestris. However, there are also host records lity represents one of the largest remaining bog­ from Quercus, Populus, Carpinus, and Betula systems in the middle of Norway. FN Nesseby: (Hansen 1956, Palm 1959). The host tree of the Rei'kejakguolba (EIS 176), 7 July 1998, leg. & species in Norway may be Betula pubescens. In col1. F0D. One male sweep-netted among Betula Sweden, the species is restricted to the south pubescens and B. nana-thickets at the margin of (Lundberg 1995), while in Denmark and in Eng­

1 5 IiJdegaard & Ligaard: Contribution to Norwegian Co/eoptera land it is widespread but rare (Hansen 1956). The than previous collectors did believe. Leiodes species probably represents a nemoral element in inordinata and Placusa suecica are typical species the Norwegian fauna, and it is considered as a detected as a result ofextensive trapping regimes. declining, monitor species in Norway (Direk­ In addition, the knowledge of the biology of rare toratet for Naturforvaltning 1999). beetle species accumulates. This knowledge makes collectors more selective in their choice of investigation areas and searching methods. There DISCUSSION is a series of examples that collectors find rare species after revealing their biology. Most species mentioned in this paper are restricted to areas in the Oslofjord-area in SE Norway. As Moreover, new species are recorded as results of pointed out in Hanssen & Hansen (1998), this area new systematic knowledge. As complex species has the highest species richness of invertebrates groups are revised and clarified, revisions ofexis­ in Norway, and many of them live in habitats of ting material often add new species to the country. restricted range such as old growth forests, sandy An example of that in this paper is Philonthus seashore habitats, large bogs, and wetlands. The pseudoparcus. Revisions ofcomplex groups may conservation value ofthese habitats in this area is also stimulate to investigations if the separated very high and many of the species and the loca­ species differ from the original ones in their bio­ lities are threatened (Hanssen & Hansen 1998, logy. This is probably the reason why we found Direktoratet for Naturforvaltning 1999). A more Hydroporus nigellus and Omalium littorale. intensive documentation of the entomological The composition of the fauna will always be a values in these localities during the last decade is process of change (Semb-Johannson 1988). a main reason why new beetle species are found Accordingly, many new Norwegian species are in Norway found as results ofrange expansions. There is good Many species are obviously overlooked by pre­ evidence that Meligethes tristis and Bruchus vious collectors for other reasons. One aspect is affinis are species whose occurrence in Norway that we are more mobile today, and hence, we are results from a northward dispersal. Many species able to visit attractive localities that earlier were living in eutrophic freshwater habitats, and hardly accessible. especially species associated with Phragmites australis and Typha latifolia, probably belong to A hilly topography causes warm microclimate in this group. These plant species have expanded south and west faced slopes in the valleys of their range and have been more abundant in SE southern Norway at summertime. Combined with Norway during the last decades as freshwater favorable soil types and scattered growth oftrees, habitats have become more eutrophic (Lid & Lid such localities may be very similar to certain areas 1994). The occurrence of Cypha suecica, Stilbus in Denmark and southern Sweden, climatically. oblongus, Psammoecus bipunctatus (in the present Many of these slopes serve as suitable habitats paper), and also Demetrias imperialis (Lindroth for thermophilous species. The records of Stenus 1973, Hansen 1994), and Alianta nigella (Hansen ochropus, Apion columbinum and A. melancho­ 1994) in Norway, therefore, may be caused by a licum are examples ofspecies that probably have recent increase in host plant or habitat range. survived in isolated populations since warmer periods in post-glacial time in Norway. Several Many species occur at the outer edge oftheir dis­ similar localities in other valleys of southern and tribution range in Norway. It can be expected that western Norway are still not satisfactory investi­ the boundary zones ofthe ranges of such species gated. Thus, there are reasons for expecting that fluctuate due to e.g. variation in climate. After more species showing disjunct distribution pat­ repeated seasons with favorable climate, some terns are present in these areas. Furthermore, new species may expand their range. Eventually, the collection methods, especially the window traps, same species may drop out if climate worsen. have shown that many species are more abundant Perhaps Dromius linearis belongs to this group

1 6 Norw. J. Entomo/. 41, 1-19. 2000 of species. Several other species reported in this Evers, Krefeld. paper are recorded from the Norwegian provinces G0nget, H. 1997. The Brentidae (Coleoptera) of adjacent to localities in our neighboring countries Northern Europe. Fauna Ent. Scand. Vol. 34. 289 pp. E.J. Brill/Scandinavian Science Press, Leiden. where the species is present. The Norwegian Hansen, M. 1988. Nogle nye danske biller (Coleoptera). records of those species may be a result of range Entomol. Medd. 56, 17-27. fluctuations. On the other hand, they may very Hansen, M. 1996. Katalog over Danmarks biller. well be overlooked previously due to their small Catalogue ofthe Coleoptera ofDenmark. Entomol. and patchy distribution areas in Norway. Random Medd. 64, 1-231. wind dispersal is probably important for the spread Hansen, M., J0rum, P., Mahler, V. & Vagtholrn-Jensen, ofthese species. Sometimes we may observe that O. 1991. Niende tillreg til «Fortegnelse over Dan­ individuals are blown far away from their area of marks bil1eD> (Coleoptera). Entomol. Medd. 59, 5­ origin. A likely example ofthat in the present pa­ 21. Hansen, M., Kristensen, S., Mahler, V. & Pedersen, 1. per is Bembidion fumigatum. 1992. 11. tillreg til «Fortegnelse over Danmarks billeD> (Coleoptera). Entomol. Medd. 60, 69-84. Acknowledgements. We would like to thank Lita Greve Hansen, M., Liljehult, H., Mahler, V. & Palm, E. 1993. Jensen (ZMB) and Lars Ove Hansen (ZMO) for the 12. til1reg til «Fortegnelse over Danmarks biller» kind loan ofmaterial. Our best thanks to Oddvar Hans­ (Coleoptera). Entomol. Medd. 61, 85-114. sen for providing material and comments on the manu­ Hansen, S. O. 1994. To nye billearter (Coleoptera) for script, Stig Lundberg for valuable information and Norge. Fauna norv. Ser B 41, 87. checking some of the identifications, Jyrki Muona for Hansen, V. 1929. Biller VII. Bladbiller og b0nnebiller verification of Mycetoporus eppelsheimians, John (Chrysomelidae og Lariidae). Danmarks Fauna 31. Skartveit for providing information and records of 401 pp. G.E.c. Gads Forlag, K0benhavn. Barvnotus moerens, TrondAndersen and the late Helge Hansen, V. 1954. Biller XVII Rovbiller 3. del. Dan­ Rinden for providing material ofOmalium littorale and marks Fauna 59.499 pp. G.E.C. Gads Forlag, K0­ Colon pseudolatum, respectively. benhavn. Hansen, V. 1956. Biller XVIII Barkbiller. Danmarks References Fauna62. 196pp. G.E.C. Gads Forlag, K0benhavn. Hansen, V. 1964. Fortegnelse over Danmarks biller Andersen,1. & Hanssen, O. 1994. Invertebrat-faunaen (Coleoptera). Entomol. Medd. 33, 1-507. pa elvebredder - et oversett element I. Biller (Coleo­ Hansen, V. 1965. Biller XXI. Snudebiller. Danmarks ptera) ved Gaula i S0r-Tnmdelag. Oppdrags­ Fauna 69.524 pp. G.E.C. Gads Forlag, K0benhavn. melding. 326, NINA Norsk institutt for naturforsk­ Hansen, V. 1973. Biller XII. Andet oplag med tillreg. ning, Trondheim. Heteromerer. Danmarks Fauna 50.307 pp. G.E.C. Ashe, 1. S. 1991. The larvae of Placusa Mannerheim Gads Forlag, K0benhavn. (Coleoptera: Staphylinidae), with notes on their Hanssen, O. & Hansen, 1. O. 1998. Verneverdige insekt­ feeding habits. Entomol. Scand. 21, 477-485. habitater Oslofjordsomn'idet. Oppdragsmelding Bakke, A. 1961. En oversikt over vart kjennskap til 546, NINA Norsk institutt for naturforskning, Norges insektfauna. Fauna (Oslo) 14,41-62. Trondheim. Bakke, A. 1999. High diversity of saproxylic beetles in Hanssen, 0., 0degaard,F. & Kvamme, T. 1997. Forslag a hemiboreal mixed forest reserve in the south of til mdliste for norske insekter. Del I. Biller (Coleo­ Norway. Scand. 1. For. Res. 14, 199-208. ptera). Fagrapport 031, NINA Norsk institutt for Direktoratet for Naturforvaltning 1999. Nasjonal naturforskning, Trondheim. mdliste for truede arter i Norge 1998. Norwegian Heijerman, T. 1993. Coeliodes nigritarsis nieuw de Red List 1998. DN-rapport 3, 1-161. Nederlandse fauna (Coleoptera: Curculionidae). Erichson, B. 1999. Nya fynd av rapsbaggar (Coleoptera: Entomol. Ber. 53, 183-184. Nitidulidae: Meligethes) med tre fOr Sverige nya Hellen, W (ed.) 1939. Catalogus Coleopterorum Daniae arter. Entomol. Tidskr. 120, 117-125. et Fennoscandinaviae, 129 pp. Helsingfors. Freude, H., Harde, K. W & Lohse, G. A. 1964. Die Johnson, C. 1966. The Fennoscandian, Danish and Bri­ Kiifer Mitteleuropas. Band 4. 264 pp. Goecke & tish species of the genus Ernobius Thomson (Col. Evers, Krefeld. Anobidae). Opusc. Entomol. 3 I, 81-92. Freude, H., Harde, K. W & Lohse, G. A. 1969. Die Johnson, C. & Lundberg, S. 1977. Placusa cribrata n. Kiifer Mitteleuropas. Band 8. 388 pp. Goecke & sp. and P suecica n. sp. from Sweden (Coleoptera:

1 7 edegaard & Ligaard: Contribution to Norwegian Coleoptera

Staphylinidae). Entomol. Scand. 8, 71-73. 104. Koch, K. 1989a. Die Kiifer Mitteleuropas. Okologie I. Nilsson, A. N. & Holmen, M. 1995. The aquatic Ade­ 440 pp. Goecke & Evers Verlag, Krefeld. phaga (Coleoptera) ofFennoscandia and Denmark. Koch, K. 1989b. Die Kiifer Mitteleuropas. Okologie 2. n. Dytiscidae. 192 pp. E.1.BriIl/Scandinavian 328 pp. Goecke & Evers Verlag, Krefeld. Science Press, Leiden. Koch, K. 1992. Die Kiifer Mitteleuropas. Okologie 3. 0k1and, K. A. 1981. Inndeling av Norge til bruk ved 440 pp. Goecke & Evers Verlag, Krefeld. biogeografiske oppgaver - et revidert Strand system. Lawrence, 1. F. & Newton, A. F. 1995. Families and Fauna (Oslo) 34, 167-178. subfamilies of Coleoptera (with selected genera, Ottesen, P. 1993. Norske insektfamilier og deres arts­ notes, references and data on family-group names). antall. NINA Utredning 055, Norsk institutt for Pp. 779-913 in Pakaluk, 1. & Slipinski, S. A. (eds.). naturforskning, Trondheim. Biology, Phylogeny, and Classification of Coleo­ Palm, E. 1996. Nordeuropas Snudebiller (Col.: Curcu­ ptera: Papers Celebrating the 80th Birthday ofRoy Iionidae). Part 1: Brachycerinae og Otiorhyncerinae. A. Crowson. Museum i Instytut Zoologii PAN, Danmarks Dyreliv 7. 356 pp. Apollo Books, Sten­ Warszawa. strup. Lid, 1. & Lid, D. T. 1994. Norsk Flora, 6. utgave. 1014 Palm, T. 1941. Zwei neue Colon-Arten aus Schwe­ pp. Det Norske Samlaget, Oslo. dischen-Lappland nebst einer Bestimmungstabelle Lindroth, C. H. 1960. Catalogus Coleopterorum Fenno­ der aus Nord-europa bisher bekannt gewordenArten scandiae et Daniae. 476 pp. Entomologiska Siill­ der Untergattung Curvimanon (Col. Silphidae). En­ skapet i Lund. tomol. Tidskr. 62, 158-165. Lindroth, C. H. 1973. Sentida f6riindringar i den Palm, T. 1948. Skalbaggar. Coleoptera Kortvingar: Fam. nordiska insektfaunaen. Exempel thin carabidema. Staphylinidae, Underfam. Micropeplinae, Olisthae­ Entomologen 2, 1-8. rinae, Proteininae, Omalinae. Svensk Insektfauna Lindroth, C. H. 1985. The Carabidae (Coleoptera) of 38. 133 pp. Entomologiska F6reningen i Stockholm, Fennoscandia and Denmark. Fauna Ent. Scand. Vol Stockholm. 15 (I). 226 pp. E.1. BrilllScandinavian Science Palm, T. 1959. Die Holz- und Rinden-Kiifer der siid­ Press, Leiden. und Mittelschwedischen Laubbiiume. Opusc. Ento­ Lindroth, C. H. 1986. The Carabidae (Coleoptera) of mol. Suppl. 16, 1-371. Fennoscandia and Denmark. Fauna Ent. Scand. Vol Palm, T. 1961. Skalbaggar. Coleoptera Kortvingar: Fam. 15 (2). 226-498 pp. E.1. Brill/Scandinavian Science Staphylinidae, Underfam. Oxytelinae, Oxyporinae, Press, Leiden. Steninae, Euasthetinae. Hiifte 2. Svensk Insektfauna Lundberg, S. 1964. Bidrag til kiinnedom om 48. 126 pp. Entomologiska F6reningen i Stockholm, skalbaggsfaunaen pi! Gotska Sand6n. Entomol. Stockholm. Tidskr. 85, 45-48. Palm, T. 1962. Bidrag til kiinnedomen om svenska skal­ Lundberg, S. 1969. Bidrag till kiinnedom om svenska baggars biologi og systematik. 42 - 47. Entomol. skalbaggar 12. Entomol. Tidskr. 90, 217-224. Tidskr. 83, 185-198. Lundberg, S. 1986. Catalogus Coleopterorum Sueciae. Palm, T. 1966. Skalbaggar. Coleoptera Kortvingar: Fam. Naturhistoriska Riksmuseet, 155 pp. Entomologis­ Staphylinidae, Underfam. Habrocerinae, Tricho­ ka F6reningen, Stockholm. phyinae, Tachyporinae. Hiifte 4. Svensk Insektfauna Lundberg, S. 1995. Catalogus Coleopterorum Sueciae. 50.93 pp. Entomologiska F6reningen i Stockholm, Naturhistoriska Riksmuseet, 224 pp. Entomologis­ Stockholm. ka F6reningen, Stockholm. Palm, T. 1968. Skalbaggar. Coleoptera Kortvingar: Fam. Miinster, T. 1923-1924. Nye fund og findesteder. B. Staphylinidae, Underfam. Aleocharinae (Deinopsis­ Coleoptera. Norsk Entomol. Tidsskr. 1,254-274. Trichomicra) Hiifte 5. Svensk Insektfauna 51. 112 Muona, 1. 1983. Two new Palaearctic Atheta species pp. Entomologiska F6reningen i Stockholm, Stock­ (Coleoptera, Staphylinidae). Ann. Entomol. Fenn. holm. 49,57-58. Palm, T. & Lundberg, S. 1993. Tilliigg till Svensk In­ Muona, 1. 1995. Taxonomic notes on the genus sektfauna, kortvingar (Coleoptera, Staphylinidae) Philhygra Mulsant & Rey (Coleoptera, Staphyli­ hiifte loch 2. Entomol. Tidskr. 114, 161-172. nidae). Entomol. Medd. 63, 11-16. Schoyen, W. M. 1879. Supplement til H. Siebke's Nilsson, A. 1994. Revision ofthe Hydroporus nigellus enumeratio. Christiania Vidensk.-selsk. Forhandl. complex (Coleoptera: Dytiscidae) including multi­ 3,11-75. variate species separation. Entomol. Scand. 25, 89­ Semb-Johannson, A. 1988. Endringer i dyrelivet i var tid. Fauna (Oslo) 41,73-90, 109-132.

18 Narw. J Enlama/. 41, 1-19. 2000

Silfverberg, H. 1979. Enumeratio Coleopterorum Fen­ Hansen, Michael. 1999. World Catalo­ noscandiae et Daniae. 79 pp. Helsingfors Entomo­ gue of Insects. Volum 2. Hydrophiloi­ logiska Bytesforening, Helsinki. Silfverberg, H. 1992. Enumeratio Coleopterorum Fen­ dea (s. str.) (Coleoptera). 416 s. Apollo noscandiae, Daniae et Baltiae. 94 pp. Helsingfors Books, Kirkeby Sand 19, DK-5771 Sten­ Entomologiska Bytesforening, Helsinki. strup, Denmark. ISBN 87-88757-31-5. Strand, A. 1939. Neue paHiarktiche Arten der Gattung DKK 690,- + porta. Ved abonnement pa Gyrophaena Mannh. (Col. Staph.). Norsk Entomol. serien gis 10% rabatt. Tidsskr. 5, 108-11 l. Strand, A. 1946. Nord-Norges Coleoptera. Tromso Mus. Arsh. 67 (1944), m.l, 1-629. Det danske forlaget Apollo Books har na utgitt Strand, A. 1957. Uber die nordischenArten der Gattung andre bind i serien med den ambisiose tittelen Liodes Latr. (Col. Liodidae). Norsk Entomol. Tids­ World Catalogue ofInsects. Denne gangen er det skr. 10, 119-130. vannkjrerene (Hydrophiloidea) som er katalogi­ Strand, A. 1965. Koleopterologiske bidrag XI. Norsk sert. Forfatteren er den svrert produktive danske Entomol. Tidsskr. 13,82-91. entomologen Michael Hansen. Strand, A. 1977. Additions and corrections to the Nor­ wegian part of Catalogus Coleopterorum Fenno­ Det er meget hyggelig at det igjen er blitt fart i scandiae et Daniae. Second series. Norw. 1. Ento­ utgivelsene av verdenskataloger pa biller. Apollo mol. 24,159-165. Books har jo na utgitt bade Hydraenidae og Hy­ Tommenis, B. A. & Breistein, 1. 1995. Fragmenterings­ drophiloidea, mens kataloger pa Ptiliidae, Crypto­ forsok i granskog. Problemstillinger og metoder phagidae og Cryptocephalinae er under bear­ samt resultater fra feltsesongen 1994. Oppdrags­ melding 342, NINA Norsk institutt for natur­ beidelse. Wroclaw, Polen har ogsa nylig utgitt ver­ forskning, Trondheim. denskataloger pa gruppene Histeridae og Cassi­ Tommenis, B. A., 0degaard, E, Breistein, 1., Wilman, dinae. I disse tider med fokus pa global biodiver­ B. & Gjershaug, J. O. 1997. Fragmenteringsforsok sitet og bevaring av artsmangfold ma det sies at i granskog Mosvikprosjektet. Oppdragsmelding grunnleggende oversikter av dette slaget er meget 488, NINA Norsk institutt for naturforskning, viktige bidrag til fremdriften innen dette enonne Trondheim. fagfeltet. Vik, A. 1991. Catalogus Coleopterorum Norvegica. 157 pp. Stig Otto Hansen, Larvik. Verdenskatalogene inneholder basisoversikter Vik, A. 1995. Latridiidae (Muggbiller). Norske insekt­ over taksonomisk status og utbredelse pa global tabeller 15. 15 pp. Norsk Entomologisk Forening, basis for den gruppen som er behandlet. En slik Oslo. katalog utgjor med andre ord et fundamentalt dokument som kan tjene til inspirasjon og enonn Received 15 November 1999, arbeidsbesparelse for alle som vii jobbe videre accepted 24 February 2000 innen gruppen enten det er snakk om systematiske, okologiske eller rent faunistiske arbeider. Katalo­ gene vii derfor vrere av interesse for museer og andre vitenskape1ige institusjoner over hele verden og er en forutsetning for entomologer som onsker seg inngaende oversikt over bestemte grupper inn­ en stone geografiske regioner. Den forrige fullstendige katalogen over Hydrophi­ loidea kom ut i 1924. Den gang inneholdt katalo­ gen 1370 kjente arter. Med Michael Hansens kata­ log er 2803 arter registrert i denne overfamilien, noe som skulle understreke hvor kjrerkommen denne katalogen er for den som onsker a jobbe med vannkjrer pa regional basis.

1 9 Book review • Bokanme/de/se

Boken har en kort og konsis introduksjon som bergs Enumeratio fra 1992. Laccobius biguttatus forklaring til katalogdelen. De nomenklaturiske Gerhardt, 1877 skal na hete L. colon (Stephens, endringene som er gj ort samt en klassifiserings­ 1829) pga. prioritetsprinsippet, dvs. at det f0rste tabell med antall arter i hvert h0yere takson (over publiserte navn skal vrere gyldig; Cercyon atri­ artsniva) finnes ogsa i innledningen, noe som er capillus (Marsham, 1802) skal hete C. nigriceps svrert nyttig. Selve katalogdelen er basert pa innga­ (Marsham, 1802) pga. f0rste revisors prinsipp, ende studier av hele 2000 vitenskapelige artikler dvs. at synonymi skal avgj0res av f0rste revisor om systematikk hos Hydrophiloidea. Katalogen hvis navnene er publisert samtidig; Megasternum bestar av en systematisk liste over alle h0yere obscurum (Marsham, 1802) skal hete M. con­ taksa og alfabetiske lister over alle gyldige arter cinnum (Marsham, 1802) pga. primrer homonymi, innen hver slekt. For alle taksa refereres original­ dvs. at da arten ble beskrevet var det allerede en beskrivelsen og for slektsnavnene refereres i til­ annen art som hadde samme navn. legg typearten og kj0nnet. Artsnavnene refererer Katalogen er ikke helt oppdatert pa utbredelse nar typelokalitet, eventuelle lectotyper eller neotyper det gjelder Norge pga. at Silfverbergs Enumeratio og referanse til f0rste bruk av ulike kombinasjoner fra 1992 er brukt som referanse for norske arter. med slektsnavn. Alle synonymer samt de viktigste Etter 1992 har tre nye vannkjrer blitt publisert fra feilstavinger er listet kronologisk. Et lite minus Norge, Helophorus fulgidicollis (Fauna norv. Ser er at det ikke er nevnt hvor typematerialet for hver B 41 :45-47), Helophorus obscurus (Fauna norv. enkelt art befinner seg. Slik informasjon hadde Ser B 45:79) og Laccobius colon (L. biguttatus) vrert til stor nytte for taksonomer som har behov (Fauna norv. Ser B 41 :45-47). Det kan imidlertid for a studere typene. Til slutt, under hvert artsnavn, ikke forventes at en katalog av et slikt omfang listes artens navrerende utbredelse pa landsbasis skal ta hensyn til nasjonale faunistiske artikler. innenfor hver biogeografiske region. Nar det Det er pa mange mater mer ryddig a forholde seg gjelder store land, som for eksempel Australia, til de regionale utbredelseskataloger som referan­ Canada, Kina, Russland og U.S.A er utbredelsen ser i denne sammenhengen. mer detaljert angitt. Etter katalogdelen finnes tre appendix som inneholder lister over I) fossile Prisen pa boka har dessverre blitt mer en det dob­ arter, 2) nomina nuda, dvs. navn som ikke refererer belte av bind 1 noe som imidlertid kan forsvares til synonymer av arter i gruppen, og 3) arter som ved at prisen per side faktisk er lavere. Mer enn opprinnelig ble beskrevet under Hydrophiloidea, 700 NKR er sannsynligvis allikevel i overkant av men som er ekskludert fra denne overfamilien. hva den generelt interesserte entomolog villegge i denne type litteratur. En slik grundig revisjon av nomenklaturen innen­ for en taksonomisk gruppe innebrerer ofte at noen I sum sa er dette et enormt omfattende arbeide av arter ma skifte navn. Mange entomologer fortviler meget h0y kvalitet og av stor vitenskapelig verdi over slike navnebytter, men dette er en n0dvendig­ som sikkert vii sta som en av de mest sentrale het for a oppna malsettingen med vitenskapelige referanser pa Hydrophiloidea i mange tiar frem­ navn, nemlig stabilitet, universalitet og unikhet. over. leg kan derfor bare gratulere Michael Han­ Tmsten far vrere at nomenklaturen tross alt sen sa mye med nok en kjempeflott innsats. All konvergerer mot det som virkelig er riktig i f01ge honn0r ogsa til Apollo Books som har tatt pa seg ICZN (International Code of Zoological Nomen­ denne krevende og meget viktige oppgave a pu­ clature), som er regelverket for zoologisk navn­ blisere verdenskataloger pa insekter. setting. Nytten og kompleksiteten av et slikt «opp­ renskningsarbeid» synliggj0res om en setter seg inn i det nomenklaturiske virvaret som f01ger man­ Frode lJdegaard ge av de vanlige artene. Det er ikke uvanlig at en art har mer enn 10 synonymer. Michael Hansens katalog innebrerer tre navneend­ ringer for norske vannkjrer i forhold til Silfver­ 20 Norw. J. Entomol. 41, 21-23. 2000

A new species of Archisotoma Linnaniemi, 1912 from the Baltic coast of Latvia (Collembola, Isotomidae)

Arne Fjellberg & Edite Jucevica

Fjellberg, A. & Jucevica, E. 2000. A new species ofArchisotoma Linnaniemi, 1912 from the Baltic coast of Latvia (Collembola, Isotomidae). Norw. 1. Entomol. 47, 21-23.

Archisotoma martae sp. n. is described from a sandy seashore habitat near Ventspils, Latvia. The small species resembles A. theae Fjellberg, 1980, but is easily identified by an exceptionally strong mucro which is half as long as the dens.

Key words: Collembola, Isotomidae, Archisotoma martae sp.n., Latvia.

Arne Fjellberg, Milgereveien 168. N-3145 Tjeme. Norway Edite Jucevica. Institute ofBiology, University ofLatvia. Miera iela 3, LV-2169 Salaspils, Latvia

INTRODUCTION DESCRIPTION The genus Archisotoma has a worldwide distribu­ Size of reproductive adults 0.5 mm. Colour dif­ tion with about 20 species in marine littoral habi­ fusely brownish gray, eyespots black. Ocelli 8+8, tats. The larger species are active on the surface of which two are smaller. PAO elongate, about 3 and may sometimes be observed in huge numbers times longer than nearest ocellus (Figure IH). on mudflats and rocky shores at low tide. The Body slender, of tubular shape, extremities short. smaller species are interstitial among sand and Mouth cone prominent, swollen. Body hairs short, gravel, and some may live in sandy beach mead­ without differentiated macrochaetae. Integument ows well above the high tide mark (Thibaud & with small sharp granules, appearing like sand­ Christian 1997). The present paper reports a new paper at high magnification. Antennae with a nor­ species which was detected among sand on the mal set of sensilla. Ant. 1 with 13 setae. Labrum Baltic coast of Latvia. with 5-5-4 setae, the two anterior rows set near apex. Frontoclypeal field with 2 prelaral setae only Archisotoma mar/ae Fjellberg & Jucevica, sp. n. (Figure 1E). Maxillary outer lobe with simple palp and 4 sublobal setae which are set close together Type material: Holotype. Female (slide) from in a bundle. Maxillae long and narrow (Figure IG). «Latvia. Ventspils. 27.VI.1998. In gravel at sea­ Capitulum is reduced to a small hook-like tooth, shore. E. Paulina leg.», deposited at Natural His­ and 5 lamellae are recognised. The longest lamella tory Museum, Department of Entomology, Lon­ bears a row of strong cilia along its dorsal edge don. Paratypes. 15+2 specimens (2 slides) together and a row ofvery thin cilia along the ventral edge, with holotype, 4+4 specimens (in 2 slides) at In­ in addition to some fine ciliation near apex. A stitute ofBiology, University ofLatvia, Salaspils. broad, almost smooth lamella is located ventrally Etymology. We dedicate our new species to Mrs. to the longest lamella. Two very short lamellae, Marta Paulina, the mother ofthe second author. one with fine marginal ciliation - the other almost smooth, are situated near the hook-like tooth. Proximally to these there is a small membraneous

21 Fje//berg & Jucevica: A new species of Archisotoma (Co//embo/a, /sotomidae) from Latvia

_.' ~ '", ..,." i i'i'" I " ~ ~ ... I' ' }('\_ - --T~::--f' >, ~ ".... /- ­ -, ~ ~o~ J ~,,_

.- \ ~ ...... F'nly'/ ...... ,\ , , t/~ ,.,-',:,t" '\'Io$r ~ h~~i' ~ IL " , , I "", '- <. r' "r," , ':' \ "'~ \" ~~-=-::- (~ ~~ 'ri~ '(A~ "I \ , , ,, , ]J",, 1 I",~" ' ... ", ~' \~: ~I-V--i" f' '"I -; , !' "\\ - )MW' ~------C\,' ~ -#A G ~ /" ...... ' I ,.. ,,. : '\ '\ '\ , '\ ~~ f ~ , ''I ,,", ""'~ I I l/" ? ?:'\ I ~" rf.'JXi\\'\~~"i~· r... I' , I " " ? ft, ~ ~ ~, ~,h~j11 r, ':'" 'I,' ...'\.. ,. '. ..,..),'~ '.' ~\1\ . ~ '\i~",==-----:1--;\'~ \. "'0°~ ~ /J/ \. r l ,. 1" :"~ ~ ... "" ... 0..... J' t t t 1 f f ~~" , " " ~ f ,,1,·'\ ",''i Qcao~ )111c11~ I';I~') ~

~.,. , r ~:-',~ ~'\"'''~' ~rft't' , r ' I t t ... " 'I ~~H f r ~ ':' 'I 'I 1,1.. ~ \1 r I' r' f; 'I ... ' '\... ,'I I A" ( t: I ~ '~ .. \ ,,~rf ~ ""',' 1.1~ . , I,,'/:'i'; ""I 11 I I 2­ r" ~lr"I\"" ! \:,/, ,I~L'U i /.,/1' \ /t ~'.!-" ~~ '1 J' ~f.'!)', ': ,\ ,t\.: n1 ''''\I. I "'\ ) ~-I,b:.,:.1~"yll H~ h2 J~

Figure 1. Archisoloma mar/ae sp. n. A. General chaetotaxy; B. Furca, dorsal view; C. Left dens and mucro, lateral view; D. Right dens and mucro, ventral view; E. Labrum and frontoclypeal field, prelabral setae encircled; F. Claw and tip of right tibiotarsus 3; G. Dorsal side of left maxilla, with alternative views of the two longest lamellae; H. Left eyefield with PAO; I. Labial hooks (hypostomal setae); J. Abdominal segments 3-6, profile.

22 Norw J Enlomo/. 47, 21-23. 2000 lamella with a few long serrations. Mandibles nor­ mal, with strong molar plate and 3-4 apical teeth. The labial palp is not analysed in detail, but ap­ pears normal for the genus (Fjellberg 1999). Proxi­ mal field with 3 setae only. Labial hooks (hypo­ stomal setae, H) strongly curved, apically flattened (Figure 11). Basomedian field of labium with 4 setae, basolateral with 5. Head with 4 postlabial setae on each side along the ventral line. Dorsal chaetotaxy as Figure lA. Head with 3-4 unpaired median setae in anterior part. Abd.I-3 with 3+3 axial setae. Sensilla hardly differentiated from or­ dinary setae, except a lateral sensillum on Th.2 and a pair of dorsomedian sensilla on Abd.4. Abd.6 bears 3+3 erect dorsomedian sensilla (Fig­ ure 1J). Ventral setae absent on thorax. Ventral tube with 4+4 distal (lateral) setae. Furca as Figure lB­ D. Dens short, with 8 dorsal setae and 4 ventral ones in apical part only. Mucro strongly devel­ oped, incurved, about half as long as dens. The inner basal tooth appears as a small hook in the middle of the dorsal edge. Lateral seta present. Retinaculum with 4+4 teeth, no seta on corpus. Tibiotarsi with 9 or lO setae in the apical whorl. Tib.l and 3 with a curved, thickened dorsoapical seta in assymmetric position. Femur of last pair of legs with an apical fleshy spur. Claws short, unarmed (Figure IF).

DISCUSSION The new species is very similar to A. theae Fjell­ berg, 1980, which differs by having only 6 dorsal setae on dens and a smaller mucro which lacks the lateral seta. The exceptionally large mucro, being halfas long as dens, immediately identifies A. martae from all other species of the genus.

References Fjellberg, A. 1999. The labial palp in Collembola. Zool. Anz. 237, 309-330. Thibaud, J.-M. & Christian. E. 1997. Biodiversity of interstitial Collembola (Insecta) in sand sediments. Eur.1. Soil Biol. 33, 123-127.

Received 3 February 2000, accepted 20 February 2000

23 Short communication

Mitostoma chrysomelas (Hermann, The species is present in many European coun­ 1804) (Opilionida, Nemastomatidae), tries, including Denmark and Sweden (Meinertz a harvestman new to Norway 1964, Roewer 1911).

Kjell Magne Olsen Acknowledgements. The juvenile specimen from 1996 was sent to Jiirgen Gruber in Austria, who kindly veri­ fied my determination. Olsen K.M. 2000. Mitostoma chrysomelas (Hermann, 1804) (Opilionida, Nemastomatidae), aharvestman new References to Norway. Norw. 1. Entomol. 47, 24. Meinertz, N.T. 1964. Eine zootopographische Unter­ suchung iiber die diinischen Opilioniden. Yid. Medd. The opilionidMitostoma chrysomelas has on two occa­ Dansk nat.-hist. For. 125,417-449. sions been collected at a locality in Engalsvik in 0stfold Olsen, K.M. 1995. Opilio parietinus (De Geer, 1778) county, Norway. Both adult andjuvenile specimens were (Arachnida, Opiliones) does belong to the Norwe­ present. gian fauna. Fauna norv. Ser. B 42, 66-67. Roewer, C.-F. 1911. Ubersicht derGenera der Subfami­ Key words: Mitostoma chrysomelas, Opilionida, Nor­ lie der Phalangiini der Opiliones Palpatores nebst way. einiger neuer Gattungen und Arten. Archiv Natur­ gesch. 77 I (Suppl. 2), 1-106. Kjell Magne Olsen, Skarvelokka, 4818 FlErvik. Norway. Solhoy, T. 1982. Trogulus tricarinatus (L., 1767) (Opili­ ones, Trogulidae) recorded for the first time in Nor­ way. Fauna norv. Ser. B 29, 48. It has now been some years since last time a new Stol, I. 1993. Check-list of North European Opiliones. harvestman was reported from Norway, viz Fauna norv. Ser. B 40, 77-79. Trogulus tricarinatus (L., 1767), discovered by Strand, E. 1900. Zur Kenntniss der Arachniden Norwe­ Solh0Y (1982). Stol (1993) proposed to exclude gens. Kg!. Norske Vidensk. Skr. 1900 (2), 1-15. Opilio parietinus (De Geer, 1778) from the Nor­ Received 15 September 1999, wegian list, but as Olsen (1995 and unpubl.) has accepted 30 November 1999 demonstrated, the species has been found on sev­ eral occasions in later years, and this leaves the statements by Strand (1900) reasonably credible. Sixteen species of harvestmen have so far been published from Norway (Stol 1993). Mitosoma chrysomelas (Hermann, 1804) has so far been collected from a single locality in Nor­ way, viz. Engalsvik, Fredrikstad, 0stfold county, UTM(wGs84) 32V NL991695 (EIS 19). One ju­ venile was collected on 18 April 1996 and two specimens (1 juv., I adult d) on 7 June 1998. The finding site is close to sea level, and strongly in­ fluenced by human activity. M. chrysomelas is easily separated from other Norwegian species: it is a member ofthe Nemasto­ matidae, with no pedipalpal claw and capitate se­ tae on the long pedipalpae, and it has silvery or golden spots on the abdomen. Adults also have rows of distinct bifid or trifid tubercles on the cephalothorax and tergites.

24 Norw. J. Enlomo/. 41, 25-28. 2000

Acid rain and the disappearance of the Apollo butterfly (Parnassius apollo (L., 1758}) from coastal areas in Norway

Geir O. Gogstad

Gogstad, G.O. 2000. Acid rain and the disappearance of the Apollo butterfly (Parnassius apollo (L., 1758)) from coastal areas in Norway. Norw. 1. Entomol. 47, 25-28.

Sedum maximum, the major host plant for lowland Parnassius apollo, was sampled from current and previous locations for the Apollo, and pH was measured after suspending the plants in de­ ionised water. The plants were generally 0.3-0.5 pH-units more acidic in current than in previous Apollo-locations. Also, the buffer capacity of the plants was measured by titration with HCI-solu­ tion. The buffer capacity increased from June to September, probably reflecting accumulation of organic material, and hence the nutritional value of the plants. The buffer capacity developed con­ siderably more throughout the summer in plants from current Apollo locations, than in plants from previous coastal Apollo locations and locations near Oslo. A correlation between buffer capacity of the host plants, and the current appearance of Apollo is thus indicated. Since the lowest pH and highest buffer capacity is found at current Apollo locations, the study indicates that other explana­ tions than acid rain should be sought to explain Apollo's retraction to inland locations. More local factors may have influenced on the nutritional value of Sedum maximum and thus caused Apollo's disappearance.

Key words: Parnassius apollo, Sedum maximum, acid rain.

Geir Gogstad, Kjetils vei 8B, N-0494 Oslo, Norway. E-mail: [email protected]

INTRODUCTION pollution, and acid rain (Bengtsson 1989), as well as climate changes, agricultural and forestry ac­ The population ofthe Apollo butterfly (Parnassius tivities/expansion, and human activities (Hansen apollo (L., 1758)) was gradually reduced from the 1993). early 1900's in the coastal regions in Southern Norway. At about 1960, but no later than 1970, The most frequently mentioned cause for the re­ the butterfly had totally disappeared from these duction of the Apollo population, is acid rain, al­ districts (Opheim 1969, Bakke 1975). Apollo has, though there also are many contradictions to this however, a stable and may be also increasing popu­ hypothesis (Hansen 1993). Acid rain may cause lation in inner districts ofNorway (Hansen 1993). liberation oftoxic aluminium from the rocks and A similar situation is found in Sweden where soil (NIVA 1998, 1999), which in turn may influ­ Apollo disappeared from most coastal locations ence on the survival of the caterpillars. We have from 1900 to 1970. (Nordstmm 1955, Janzon and measured the pH in Apollo's host plant Sedum Bignert 1979, Henriksen & Kreutzer 1982). One maximum from various districts of southern Nor­ remarkable exception from Sweden is the frequent way throughout 1998 and 1999 in order to inves­ appearance ofApollo at Gotland. tigate this possibility. Numerous causes for the disappearance ofApollo have been suggested: parasites, infectious agents, MATERIALS AND METHODS cat attacks, traffic related destruction, pesticides, Sedum maximum, was sampled from previous and butterfly collectors, lack of host plants, chemical currentApollo locations in August 1998, and from

25 Gogstad: Acid rain and disappearance of the Apollo butterfly (Parnassius apollo) in Norway

June to September 1999. The coastal collection from plants sampled at the same location on the sites where Apollo have been previously reported same date showed negligible variations (0.03 pH were Sandefjord (VE) (appearance of Apollo as units or less) and are not listed separately. The late as 1959 (Sverre Sandberg, personal commu­ values obtained with plants from the two coastal nication)), and Engelsviken in Fredrikstad com­ locations and from Maridalen, Oslo, all demon­ munity, (0) (assumed previous location). Host strated higher pH than in the plants from the in­ plants were also collected from Maridalen, Oslo land location of Flit and Nesbyen, and from (AK) which is assumed to be a previous location, Kviteseid. Thus, plants from current Apollo loca­ since Apollo was regarded as common in the Oslo tions were generally more acidic than plants from district until about 1920 (Hansen 1993). Collec­ non-Apollo-locations. The values obtained from tions were also made from Begnadalen in Slilr­ Slilr-Aurdal resembled the values from the other Aurdal community (OS), which is another previ­ inland locations. The pH showed some variations ous inland Apollo location and assumed to still over time (June-Sept) in the year 1999. However, have populations (Hansen 1993). Examinations the general tendency throughout the year was that from Begnadalen to Bagn inAugust 1998 and 1999 the pH was 0.3 to 0.5 units lower at current Apollo­ did not, however, result in any observations. The locations. collection sites from current Apollo locations were To investigate the plant's buffer capacity, the sus­ Flit and Nesbyen in Hallingdal (BV), and one col­ pensions were titrated with 0.1 mol/L HCI. The lection from Kviteseid (TEI) was also made. results are shown in Table 2. Each collection included three different plants at The plants collected in June demonstrated a mark­ each date and location. All samples were kept fro­ edly lower buffer capacity than plants collected zen at -20°C until analysed. later in the summer. It is also seen that plants from Analyses were performed by adding 10.0 g ofleafs the current Apollo locations in Flit and Nesbyen, to 200 mL of de-ionised water produced by a and Kviteseid, as well as from the inland location MilliQ system. The water had no detectable buffer in Slilr-Aurdal, developed a better buffer capacity capacity upon addition of0.1 mollLHCI. The leafs during the summer than the plants from the other were suspended in the water using a Warren locations. The plants from Maridalen, Oslo, dem­ Blendor until a homogenous mixture with no ma­ onstrated the lowest buffer capacity. The plants jor particles or fibres were left. from Sandefjord demonstrated values in between those from inland and from Maridalen, Oslo. The pH was measured using a Hanna HI 8417 pH­ meter, after adjustment with standard buffer solu­ A further observation was that some of the plant tions of pH 4.00 and 7.00. suspensions changed from a green to a strikingly red colour when titrated below a pH of about 3.0. The suspensions were also examined for buffer There is, however, not possible to see any relation capacity by titration with 1.0 mL portions of 0.1 with this colour shift and collection date, pH, mollL HCI. Plant material is expected to buffer buffer capacity, or the presence of Apollo. added H+-ions. The pH-values will therefore dif­ fer from the values obtained when the same amount of acid is added to water. The differences DISCUSSION between pH in plant suspension added acid, and This study shows that the pH in Sedum maximum the expected pH if the same amount of HCI was is more acidic at current Apollo locations than at added to a non-buffering system, were collected locations from which Apollo has disappeared. as a measure of buffer capacity. The study also indicate a relation with respect to buffer capacity in Sedum maximum, and the pres­ RESULTS ence of Apollo, in various districts investigated. Table I shows the pH in suspensions of Sedum Since the buffering substances in biological ma­ maximum from various locations. The pH values terial are dominated by proteins, phospholipids,

26 Norw J Enlomo/. 47, 25-28. 2000

Table 1. pH in suspensions of Sedum maximum collected from various locations in Southern Norway 1998-1999. Each value is the average from three individual plants.

Period Hallingdal Sandefjord Maridalen Fredrikstad SliJr-Aurdal Kviteseid Aug. 1998 5.51 5.85 6.18 5.81 5.43 Jun. 1999 5.65 5.93 5.66 5.45 Jul. 1999 5.55 5.82 5.88 5.34 5.45 Aug. 1999 5.48 5.78 5.92 5.85 5.28 Sept. 1999 5.53 5.83 6.08 5.38 Average 5.54 5.82 5.94 5.83 5.36 5.45

Table 2. Buffer capacity in suspensions of Sedum maximum collected from various locations in Southern Norway. Buffer capacity is measured as the difference in pH of 200 mL plant suspensions containing 10g leaf material, after addition of 1 mL 1 mol/L HCI, and the value calculated if the same amount of HCI was added to a non-buffering system.

Period Hallingdal Sandefjord Maridalen Fredrikstad Ser -Aurdal Kviteseid Aug. 1998 0.76 0.78 0.49 0.44 0.82 Jun. 1999 0.41 0.38 0.15 0.39 Jul.1999 1.05 0.65 0.23 0.97 1.02 Aug. 1999 1.33 0.82 0.34 0.41 1.17 Sept. 1999 1.45 1.14 0.38 1.32

and certain glycans, the buffer capacity also re­ influence on Apollo caterpillars since the pH is flects the contents of nutrients per volume of the generally lower in districts were Apollo is currently plant leafs. Since the plants' buffer capacity were found. The variations in observed pH is at the level found to increase throughout the year, this appear of 0.3-0.5 pH units, which in a biological context to reflect an accumulation of organic substances. is a quite large difference. However, the variation in the plants' buffer capacity, and thus the plants' The pH in the plants is reflected by three factors: contents oforganic material correlates to the pres­ 1) the pH in the precipitation, 2) the buffer capac­ ence of Apollo, and may be the factor critical to ity ofthe ground, and 3) the buffer capacity ofthe survival ofApollo caterpillars. plants. Acid rain will liberate toxic aluminium ions which may be absorbed by the plants. The expo­ Some factor may have influenced on Sedum maxi­ sure of host plants to aluminium ions will be di­ mum sability to accumulate organic material, and + • recdy dependent on the amount ofH -Ions exposed this factor may be the basic cause ofApollo's re­ to the ground, and the ground's buffer capacity. traction from coastal districts ofNorway. This fac­ tor has apparently not influenced on the plants' Direct measurement of host plant pH does not survival since they are flourishing and widespread alone justifY conclusions related to various levels at all the location investigated, as well as over large of aluminium toxicity. However, since the most parts of the southern Norway. acidic plants also show the highest buffer capac­ ity, this total picture indicates that they have been Acid rain is a decreasing, but still quite abundant exposed to more H+-ions, and hence more toxic environmental problem. (NIVA 1998, 1999). This aluminium ions than the plants with higher pH. study can not totally exclude that acid rain is the long tenn cause for the effects on Sedum maxi­ A lower pH does not in itself seem to have any

27 Gogstad: Acid rain and disappearance of the Apollo butterfly (Parnassius apollo) in Norway mum. Conditions in previous periods may as well mum may regain its potential to serve as qualified have been different from the short observation food for the Apollo caterpillars. period in this study. Studies on distribution ofacid rain in Norway also show that the coastal regions References ofSouthern Norway are most heavily exposed, and Bakke, A. 1975. Dagsommerfug1er. 144 pp. Cappe1en, that acidity in rain gradually decreases in the north­ Oslo. ern direction (NIVA 1998, 1999). It is, however, Bengtson, H. 1989. Apollo 6kar, sakte men slikert. Sve­ remarkable that Apollo also has disappeared in riges Natur 4, 5-6. coastal regions with calcified, buffering ground Hansen, L.O. 1993. Status of the butterflies Apollo (Hansen 1993), and that Apollo is frequently found (Parnassius apollo) and Scarce Heath (Coenonym­ at Gotland which suffers hard from acid rain from pha hero) in Norway. NINA Utrening 046, 43pp. eastern Europe. These observations together with Henriksen, H.J. & Kreutzer, I. 1982. Skandinaviens dag­ sommerfugle i naturen. 215 pp. Skandinavisk Bog­ the present study thus leaves us with an apparent forlag, Odense. negative conclusion to the theory that acid rain is Janzon, L.-A. & BignertA. 1979. Apollofjiiriln i Sverige. the major cause ofApollo's disappearance. Fauna Flora 74, 57-66. What other potential causes should be sought? As Langer, T.W 1963. Sommerfug1eliv. 126 pp. Munks­ pointed out by Hansen (1993), it is less likely that gaard, Copenhagen. NIVA. 1998. Skje1kviHe, B.L.(ed.). NIVA Report 7481 traffic, butterfly collectors, or agricultural or for­ 98. 216pp. estry activities causing fauna changes, are of any NIVA. 1999. Skjelkviile, B.L. (ed.). NIVA Report 7701 significance. We certainly agree to Hansen's ar­ 99.65 pp. guments. It may be added that iftraffic death was Nordstrmn, F. 1955. De Fennoskandiask Dagfjiiri1namas a major cause, it should be more abundant at the utbredning. Lunds Universitets Arsskrift . N.F.Avd.2 current inland locations with heavy summer traf­ Bd.51, Nr.l. C.W.K.G1eerup, Lund. fic at the roads, and closely located railways, than Opheim, M. 1969. Distribusjonsstudier av norske Lepi­ in the far less trafficked coastal locations. It has dopter 1. Atlanta Nor. 1, 121-12. also been described that Apollo caterpillars prefer Received 4 October 1999, the reddish version of Sedum maximum, and that accepted 15 February 2000 this could be linked to a critical factor for Apol­ lo's survival (Langer 1963). Our studies did not, however reveal any correlation between red col­ oured plant and pH, nor between red coloured plants and buffer capacity. It is striking that all current Apollo locations are more scarcely populated than the locations from which Apollo has disappeared. It is also striking that Apollo seems to have disappeared in parallel with development ofmodem industry and oil heat­ ing systems. The lowest buffer capacity we found in Sedum maximum was from Maridalen, close to Oslo, and from Engelsviken, close to the cities in 0stfold. Both places have high degree of pollu­ tion from industry and other human activities. Thus, other pollutants having a more local effect may also be involved in the disappearance of Apollo. Since air and water pollutants have been reduced considerably over the last decade (NIVA 1998, 1999), it is possible that coastal Sedum maxi­

28 Norw. J. En/omol. 41, 29-40. 2000

Saproxylic beetles (Coleoptera) of Troms and western Finnmark, northern Norway with exceptional distribution in Fennoscandia

Johan Andersen, Stefan Olberg and Lisbeth Haugen

Andersen, 1., Olberg, S. & Haugen, L. 2000. Saproxylic beetles (Coleoptera) ofTroms and western Finnmark, northern Norway with exceptional distribution in Fennoscandia. Norw. 1. Entomol. 47, 29-40.

Saproxylic beetle species in Troms and western Finnmark generally have a continuous distribution in Fennoscandia or their occurrences are more closely connected with the distribution in Finland and/or Sweden than in the rest of Norway. Six species constitute notable exceptions to this general pattern. Among these Scaphisoma boleti, Enicmus lundbladi and Corticaria pineti are southern or eastern species with isolated, northern occurrences in continental parts ofTroms and western Finnmark (Dividalen, MaIselvdalen, Alta). Orthocisftstivus is more or less continuously distributed through­ out Norway north to Troms, whereas Hapalarea vilis and H. ioptera are distributed in coastal areas of the country. These three species have a southern distribution or are absent in Sweden and/or Finland, except for a find of H. ioptera in northern Finland. The reasons for these distributional patterns are discussed. C. pineti is reported for the first time from Norway.

Key words: saproxylic beetles, Coleoptera, distribution in Troms and Finnmark.

Johan Andersen, Stefan Olberg and Lisbeth Haugen, Department of Ecology/Zoology, IB, Univer­ sity ofTromsl'J, N-9037 Tromsl'J, Norway.

INTRODUCTION them also have a more or less continuous distri­ bution within the area. Approximately 260 obligatorily saproxylic beetle species (live beneath bark, in wood, or in tree-in­ However, although most of the saproxylic beetle habiting fungi of debilitated or dead trees) have species of Troms and western Finnmark also are been recorded in Troms and western Finnmark in present in South Norway, relatively few species northern Norway (Lindroth 1960, Strand 1946, have been recorded in Nordland county, and sev­ 1970,1977, Vik 1995, Huse 1997, Haugen 1999, eral have not been found in the counties ofTnmde­ S. Olberg & 1. Andersen unpublished data). Nearly lag (ST, NT) (Huse 1997). These apparent dis­ all of them are confined to, or may use birch junctions in Norway may be due to insufficient (Betula pubescens Ehr., incI. B. tortuosa (Ledeb.) investigations in the areas of concern, but for a Nyman) and/or pine (Pinus sylvestris L.) as hosts number of species the break is thought to be real directly or indirectly (via tree-inhabiting fungi). (Huse 1997). Birch invaded Scandinavia soon after the Weichsel Most of the current species have a much more glaciation and was later followed by pine. Pine continuous distribution in Finland and/or Sweden. colonized Troms and western Finnmark from the The occurrences in Troms and western Finnmark S and SE, birch also from the E and NE (Gjrerevoll seem often to be more closely connected with those 1992). These tree species are now distributed in Lule Lapmark (Lu) and/or Tome Lapmark (To) through most of Fennoscandia (Hultl:ll 1971, Lid in Sweden or Lapponia Enontekiensis (Le) in Fin­ 1985) and many ofthe beetle species confined to land as is the case for Enicmus rugosus (Herbst)

29 Andersen et a/.: Saproxylic beetles (Coleoptera) from northern Norway

(Figure 1). It is likely that such species invaded Troms and western Finnmark from Finland and! or Sweden. However, some ofthe saproxylic beetle species present in Troms and western Finnmark have a distribution in Fennoscandia which devi­ ates remarkably from the usual pattern. These ca­ ses are presented and discussed in the present pa­ per which is part of a work dealing with the ecol­ ogy and distribution ofsaproxylic beetles in north­ ern Norway.

METHODS Information about the distribution of the species was taken from Lysholm (1937), Strand (1946, 1953, 1961, 1977), Hansen (1964), Lindroth (1960), Johnson (1967), Lucht (1987), Bily & Mehl (1989), Vik (1995) and Lundberg (1995). O. Hanssen, S. Ligaard, H. Rinden and F. 0de­ gaard provided additional, unpublished informa­ tion concerning the occurrence of species in vari­ ous parts of Norway whereas H. Silfverberg gave us information about their distribution in Finland. Our own investigations (reported with collector) Figure 1. The distribution of Enicmus rugosus in have mainly been made at severallocalities in TRI Fennoscandia. Specific localities are only mapped Dividalen (EIS 147) and around TRY Troms0 (EIS for northern Norway: TRI Skibotn (Leg. S, Olberg); 162, 163) using various collecting methods, e. g. F0 0vre Pasvik (Leg. S. Olberg & J. Andersen). catching with window traps and billet traps, rear­ ing from sporocarps of fungi (Huse 1997, Brattli et al. 1998, Fossli & Andersen 1998), picking by hand and sieving. These investigations were con­ ducted in 1994-1999. In May and June 1999, a

Figure 2. Map of Troms showing the localities where the occurrence of the fungus S/ereum rugosum was inves­ tigated. 1. Oldervikdalen; 2. T0nsvi­ ka; 3. Hak0ybotn; 4. Andersdal; 5. Breivikeidet; 6, Svartnes; 7. Mid­ dagsbukt; 8. Laksvatn; 9. Nordkjos­ botn; 10. Skibotn; 11. Rundhaug; 12. Dividalen, loc. 1; 13. Dividalen, loc. 2; 14. Dividalen, loc. 3.

30 Norw. J Entomol. 41, 29-40. 2000 special survey ofthe occurrence ofthe fungus Ste­ THE SPECIES reum rugosum (Pers.: Fr.) Fr. (Corticiaceae), which Hapa/araea vi/is (Erichson) is a host ofOrthocisfestivus (Gyllenhal), was made at a number of localities in Troms county from Distribution in Fennoscandia (Figure 3): Distri­ the coast in the NW of the county to the inland bution in Troms: TRY Makeskjrer (EIS 170); areas in SE (Figure 2). All dead, standing or lying Tromsfl (EIS 162); Tflnsvika (EIS 162), leg. J. An­ logs (> 10 cm in diameter at the base) of Alnus dersen, 01dervikdalen (EIS 163), leg. J. Andersen. incana (L.) Moench and Betula pubescens were Nearest records in Norway: NNV Melbu (EIS examined for occurrence of this fungus species. 137), Bfl (EIS 143) and NSI Mosjflen (EIS 118). The size of the areas varied, but at least 50 dead Nearest record in Sweden: Sodermanland (So) logs ofA. incana were examined at each locality. (Lundberg 1995). Not reported from Finland At places where S. rugosum had not previously (Lundberg 1995). Widely distributed, but rare in been looked for, samples were collected and ex­ Denmark (Hansen 1964). Total distribution: Large amined for beetles in the laboratory according to parts of Europe eastwards to Poland, Tunisia, Al­ the procedure of Fossli & Andersen (1998). geria, Asia Minor, Tbilisi, Yerevan (Strand 1961, Lucht 1987). Investigations were made at Frihetsli in Dividalen in May, June and October 1999 to unveil the habi­ Ecology. The individuals from Oldervikdalen were tat (host) selection of Enicmus lundbladi Palm. collected in window traps whereas the two speci­ Stems of dead, standing or lying logs of decidu­ mens from Tflnsvika were found under the bark ous tree species (B. pubescens, A. incana, Salix of a standing, rotten stem of Alnus incana. Al­ spp., Sorbus aucuparia L.) - with or without vis­ though Koch (1989a) reported H. vilis from com­ ible infections of various types of fungi - were post and in burrows ofrodents, the species seems examined. The bark, sometimes also the wood, was to be obligate saproxylic, at least in Scandinavia loosened and thoroughly sifted through Reitter sieves (ca. 5.5 mm mesh size). The fine fractions of the sifted material were examined in the labo­ ratory under 5 or 10 X magnification. Formost ofthe species we have only mapped spe­ cific localities for the counties of Tmndelag and northern Norway. The province division is accord­ ing to 0kland (1981) whereas the nomenclature of the species follows Lundberg (1995). New records according to the district division of0kland (1981) are marked with an asterisk. Some species present in Troms have their northernmost known occurrences in Sweden in the district Norrbotten (Nb). These species may, in fact, have rather iso­ lated occurrences in Troms, but since it is unknown to us where they have been found in the very large district Nb (between lat. 65 and 68.5 0 N) we have not treated these species in further detail. We have omitted introduced species and species that are doubtfully established in the areas ofcon­ cern.

Figure 3. The distribution of Hapa/area vllis in Fen­ noscandia. The map gives specific localities.

3 1 Andersen et al.: Saproxylic beetles (Coleoptera) from northern Norway

(Strand 1946). It is found beneath bark or moss of trees as well as conifers, and on tree-inhabiting deciduous trees as well as conifers (Hansen 1964, fungi. H. ioptera also visits flowers (Hansen 1964). Koch 1989a), according to Strand (1946) also be­ neath bark of bushes. Scaphisoma boleti (Panzer) Distributon in Fennoscandia (Figure 5): Distribu­ H. ioptera (Stephens) tion in northern Norway: TRI Rundhaug (EIS Distribution in Fennoscandia (Figure 4): Distri­ 154). FV Mattisdalen, Alta (EIS 173), leg. H. Rin­ bution in Troms: TRY Oksfjorddal (172); *TRI den. Nearest record in Norway: STI. Nearest Svartnes (EIS 162), leg. 1. Andersen. Nearest record in Sweden: Angermanland (Ang) (Lund­ record in Norway: NNV Melbu (EIS 137). Near­ berg 1995). Nearest record in Finland: Ostrobott­ est record in Sweden: Jiimtland (Jii) (Lundberg nia kajanensis (Ok) (Silfverberg in lit.). Distrib­ 1995). Nearest record in Finland: Lapponia ina­ uted throughout Denmark (Hansen 1964). rensis (Li). Also recorded in the district Lps which Ecology. Although Koch (1989a) mentioned an now belongs to Russia (Lindroth 1960). Distrib­ occurrence in Sphagnum, information from Scan­ uted throughout Denmark (Hansen 1964). dinavia indicates that the species is obligate sapro­ Ecology. A single specimen was collected under xylic (Strand 1946, Palm 1959, Hansen 1968). As the bark ofa rotten, standing stem ofAlnus incana the other members ofthe family, S. boleti is myce­ infected with Stereum rugosum at Svartnes. It was tophagous, but it is probably catholic as regards found together with Orthocis festivus. According host selection (Palm 1959). to Strand (1946), Palm (1959) and Koch (1989a) the species lives beneath the bark of deciduous

Figure 4. The distribution of Hapalarea ioptera in Fen­ Figure 5. The distribution of Scaphisoma boleti in noscandia. Specific localities are only mapped for SflJr­ Fennoscandia. Specific localities are only mapped TrflJndelag, Nord·TrflJndelag and Northern Norway. for MflJre, SflJr- TrflJndelag and Troms.

32 Norw. J. Enlomo/. 47, 29-40. 2000

Enicmus /undb/adi Palm logs infected with various species of fungi such Distribution in Fennoscandia (Figure 6): Distri­ as Stereum hirsutum (Willd.: Fr.) Gray, Plicatura bution in northern Norway: *TRI Dividalen, nivea (Fr.) Karst., several species ofPolyporaceae Frihetsli and 3 km N of Frihetsli (EIS 147), leg. and mycelia of unidentified species. L. Haugen and 1. Andersen. In Norway otherwise According to Lundberg (1964, 1969) E. lundbladi only known from VE Tj0me (EIS 19). Nearest lives beneath the bark of aspen (Populus tremula record in Sweden: Ang (Lundberg 1995). Only L.) infected with the imperfect stage of the fun­ recorded in southernmost part ofFinland: Nylandia gus Hypoxylon mammatum (Wahlenb.) 1. H. Mill. (N) and Tavastia australis (Ta) (Silfverberg in lit.). (= Entoleuca mammatum (Wahlenb.) 1. D. Rogers The species has not been reported from other & Y.- M. Ju) (Ascomycetes, Xylariaceae). How­ neighbouring countries, nor in Central Europe. ever, aspen was not observed at the localities of Ecology. Fourteen specimens were collected in concern in Dividalen and the imperfect stage of window traps in Dividalen. A majority of them H. mammatum has not yet been found in Norway were found at Frihetsli. The locality is situated in (A. Granmo pers. comm.). Imperfect stages of a SW-W-facing slope in the valley. The area is other more or less closely related taxa occur on forested with Alnus incana, Betula pubescens, various species ofdeciduous trees in northern Nor­ Salix spp., and Sorbus aucuparia. The soil at the way (A. Granmo pers. comm.) so there is a possi­ site was moist or wet and had a luxuriant herba­ bility that one of them may be the host of E. ceous vegetation. The forest contained much dead lundbladi in Troms. However, despite our repeated investigations in Dividalen, we have not been able to find E. lundbladi associated with its host. In any event, it is quite obvious that the host selec­ tion of the species is different in Troms than fur­ ther south. We find it unlikely, however, that the

Figure 6. The distribution of Enicmus lundb/adi in Fennoscandia. Specific localities are mapped for Norway. The approximate -11 0 C isotherme for January (period 1961-90) in Sweden is drawn, based on Alexandersson et al. (1996). For winter Figure 7. The distribution of Corticaria pineti in Fen­ temperatures of Troms and Finmark see Table 2. noscandia. Specific locality is mapped for Norway.

33 Andersen et a/.: Saproxylic beetles (Coleoptera) from nor/hem Norway species should be confined to a single fungus spe­ colus ater (L.). The rather open forest at the site cies with an isolated occurrence in Troms. was of a primeval type with many fallen logs of pine and some dead, standing or fallen logs of Corficaria pineti Lohse birch. The locality is situated in a SW-facing slope Distribution in Fennoscandia (Figure 7): Distribu­ in the valley. According to Koch (1989b) C. pineti tion in northern Norway: *TRI Dividalen, 3 km lives on twigs, in heaps ofbark etc. ofpine, which N of Frihetsli (EIS 147), 18 August 1997, 1 (J, suggests that the species is independent of A. leg. 1. Andersen. Not recorded elsewhere in Nor­ xantha, at least in Central Europe. way. Distribution in Sweden: Halland (Ha), Sma­ land (Srn) (Lundberg 1995). Distribution in Fin­ Orfhocis festivus (Gyllenhal) land: Nylandia: Helsinki and a few places in the north (ObN: Rovaniemi?, details unpublished) Distribution in Fennoscandia (Figure 8): 0. festi­ (Silfverberg in lit.). Not reported from Denmark. vus has neither been found in the coastal districts In Central Europe reported from Germany and nor in the most continental parts ofTroms, whereas Austria. it is common in the central fjord districts of the county: *TRI Four localities in Balsfjord (Mid­ Ecology. The specimen collected in Dividalen was dagsbukt, Sandoyra and Lakselvbukt (EIS 154); found by sifting old, dry wood of trunks of pine Svartnes (EIS 162), leg. S. Olberg & 1. Andersen on the ground, infected with the resupinate poly­ for all localities. Skibotn (EIS 155), leg. 1. Ander­ pore Antrodia xantha. (Fr.) Ryvarden. C. pineti was sen. Nearest records in Norway: NNV Sortlands­ found together with numerous specimens ofRhyn- eidet (EIS 144), leg. 1. Andersen. Erikstadfjord (EIS 138), Bo (EIS 143): *NN0 Skjomen (EIS 139,140, leg. T. E. Fossli. Nearest records in Swe­ den: Ang and Asele Lapmark (As) (Lundberg 1995). Nearest record in Finland: Savonia borealis (Sb) (Lindroth 1960). Distributed throughout Den­ mark (Hansen1964). Ecology. In Nordland and Troms the species has been found along brooks, small rivers or on S­ facing slopes in forests dominated byAlnus incana and Salix spp. It has been collected exclusively in Stereum rugosum on Alnus incana at the locali­ ties in Balsfjord, B0 and Skjomen. In South Nor­ way (AK Oslo) we have found 0. festivus scat­ tered in Stereum hirsutum on birch and in S. san­ guinolentum (Alb. et Schw. ex Fr.) Fr. on spruce (Picea abies (L.) Karst.) although the species was most abundant in S. rugosum. There was no indica­ tion of reproduction (occurrence of larvae, pupae or tenerals) in S. hirsutum or S. sanguinolentum whereas larvae have frequently been found in S. rugosum. It is likely, therefore, that S. rugosum is the preferred host as stated by Koch (1989b); see Figure 8. The distribution of Orlhocis festivlJs in also Hansen (1964). Table 1 gives the results of Fennoscandia. Specific localities are only mapped registrations of Stereum rugosum at a number of for Central (MRI, STI, NTI) and northern Norway. localities in Troms county from NW to SE. The The lines show the approximate northern (Finland, data indicate that S. rugosum is much more com­ Sweden) and eastern (Norway) limit for common mon on Alnus than on Betula within the area. Fur­ occurrence of AlnlJS incana (Hulten 1971).

34 Norw J. Entomo! 41, 29-40. 2000 thennore, whereas the fungus is quite common at Since parts of Norway are insufficiently investi­ the coast and in central fjord districts, it occurs gated it is likely that the rest of the species may very scatterly in the innennost parts of the fjords also be found in other districts in the country. How­ and in the valleys of the county. ever, Sweden and parts ofnorthern Finland are so well investigated that we regard the distributional DISCUSSION gaps in S-SE to be reliable. Reliability of the distribution pattern Causes of the distribution pattern of the There is a seemingly weak break in the distribu­ species tion of Orthocis festivus between Troms/norther­ most Nordland and the occurrences further south Species mainly with western distributions in Fennoscandia (Figure 8). It is likely, however, The fennoscandian distribution of Orthocis festi­ that the species is more continuously distributed vus is probably linked to the presence of Alnus throughout Norway since Nordland is incom­ incana, the host of Stereum rugosum in which the pletely investigated. The present known distribu­ beetle lives. Our investigations indicate that S. tion of Hapalarea vilis in Fennoscandia is very rugosum has a strong preference for A. incana in scattered (Figure 3), but it is probably more con­ Troms (Table 1), and, since this also is the case in tinuous since coastal parts ofNorway are incom­ northern Sweden (Strid 1975, Ryman & Holmasen pletely investigated. The species is supposed to 1992) it is probable that it applies to northern be limited to the west- and south coast ofNorway, Fennoscandia in general. Solheim (1979) reported SE Norway and the southemmost parts of Swe­ S. rugosum to be common in Dividalen. Our data den. It is also likely that H ioptera is more con­ (Table 1), as well as those ofIversen (1997) indi­ tinuously distributed in western parts of Norway cate, however, that S. rugosum is rare or absent in for the same reason. continental parts (innennost fjord districts and the valleys) ofTroms although A. incana is common

Table 1. Per cent logs of Alnus incana and Betula pubescens infected with Stereum rugosum (8) along a transect from the coast to inland in Trams county (see Figure 2). Occurrence of Orthocis festivus (0) on S. rugosum (per cent logs with infection of 0. festivus of total number infected with S rugosum) and of Dryocoetes alni (D.a.) on A. incana is also given. N= no. of logs.

Alnus incana Presence Betula pubescens Areas Locality N %(8) %(0) of D.a. N %(8) %(0) Coast 1. Oldervikdalen 145 9.7 0 + 68 2.9 0 2. Tl'Jnsvika 98 14.3 0 + 3. HAkl'Jybotn 56 16.1 0 + 15 0 4. Andersdal 80 17.5 0 + 3 0 5. Breivikeidet 78 14.1 0 ? 73 0 Fjord 6.8vartnes 95 14.7 + + 3 0 7. Middagsbukt 139 18.7 84.6* + 30 0 8. Laksvatn 136 7.4 100 ? 9. Nordkjosbotn 100 2.0 0 ? 10.8kibotn 139 1.4 0 + Inland 11. Rundhaug 113 0.9 0 + 12. Dividalen loc.1 162 0 + 44 0 13. Dividalen loc.2 52 0 + 14. Dividalen loc.3 107 0.9 0 + Mean 1,3,4,5,7,9 14.8** 1.0**

• Data from Fossli & Andersen (1998) •• Difference in infection rate significant (p < 0.01; X\I) = 10.55)

35 Andersen et af: Saproxy/ic beetles (Coleoptera) from northern Norway within the same areas. In accordance with which, among others, are characterized by low Eriksson et a1. (1984) and Ryman & Holmasen winter temperatures (see e. g. Fig. II in Dahl (1992) we therefore conclude that S. rugosum 1998). There are indications that poikilothermic must be rare in the continental parts of northern­ animals may be excluded from continental areas most Sweden and Finland especially because A. due to low temperatures during the coldest part incana is weakly represented or nearly absent of the year (Andersen & Halvorsen 1984, within a large part of that area (Figure 8). Since it 0degaard 1994). Potential habitats (beneath bark is uncertain whether O. festivus is able to estab­ of standing or fallen trees) ofH. vi/is in continen­ lish successfully in species of Corticiaceae other tal areas would frequently be uncovered by snow than S. rugosum, there is a possibility that the dis­ because oflow precipitation in the winter months tributional gap of the beetle in continental parts (F0rland 1993). Consequently, such habitats ofnorthern Fennoscandia in part is due to the scar­ would have temperatures close to those ofthe air, city or absence of a suitable host there. although temperature conditions in dense stands are dampened in comparison with those on open Other beetles (Epuraea longiclavis Sj6berg, Dryo­ ground (Munn 1966). It is therefore possible that coetes alni (Georg» as well as fungi (Plicatura H. vi/is is excluded from continental areas due to nivea) which are associated with A. incana (Le­ very low winter temperatures. kander et a1. 1977, Eriksson et a1. 1981, Koch 1989b, Ryman & Holmasen 1992, Audisio 1993) H. vi/is most probably immigrated to Scandina­ also have distributional gaps or are scatterly dis­ via from the S and Sw, as did several other spe­ tributed in northern Sweden. As for 0. festivus, cies previously regarded as Weichsel Glaciation this may be explained by absence or scarcity of survivors (see Coope 1969), and spread along the their host. coast of Norway. The distribution of Hapalarea vi/is, H. ioptera, The distribution pattern of H. ioptera in Fenno­ Scaphisoma boleti, Enicmus lundbladi, and scandia is enigmatic. The species avoids areas with Corticaria pineti can not be explained by means a markedly continental climate in most of its oftheir host selection since they either are poorly range. However, the occurrence of the species in host specific or the host (Pin us sylvestris) of the the districts Li and Lps may break this pattern. species is distributed throughout most of Although generally low, winter temperatures vary Fennoscandia. Some reservations may be taken from locality to locality within these districts, so for E. lundbladi since its host in Troms is un­ it is uncertain whether the absence of the species known. further south in Finland and Sweden is due to too low winter temperatures. Strand (1946) and Lindroth (1949) proposed that H. vi/is survived the Weichsel Glaciation on refu­ gia in western Norway. However, ice-free areas Southern or southeastern species had an arctic climate during the Weichsel maxi­ S. boleti, E. lundbladi, C. pineti and M linearis mum and were consequently uninhabitable for probably immigrated to Troms and western trees (Vorren et al. 1988, Aim & Birks 1991, Dahl Finnmark from Sweden and/or Finland. Like these 1992). Since H. vilis mainly lives beneath bark of species, the southern, tree-inhabiting fungi trees, it is unlikely that it survived the last glacia­ Ceriporia purpurea (Fr.) Donk, Dichomitus tion on refugia in western Norway. Contrary to campestris (Qu6l.) Dom. & Orlicz, Stereum the opinion of Strand (1946) and Lindroth (1949) subtomentosum Pouzar and Steccherinum we find that H. vilis has a subatlantic distribution ochraceum (Pers.: Fr.) S. F. Gray also occur iso­ matching that of some species of plants (e. g. lated in the valleys and/or fjords ofTroms and/or Carex pulicaris L.) and terrestrial snail species Finnmark (Ryvarden 1971, 1976, Eriksson et a1. (e. g. Arion ater (L.» (Dah1 1950, Hulten 1971, 1981,1984, Solheim 1979, Olofsson 1996, Iversen Kerney & Cameron 1979). H. vi/is is absent in 1997). Thus, isolated occurrences of southern or continental parts ofEurope (E and NE ofPoland) southeastern species in northernmost Norway may

36 Norw J Enlomo/. 41, 29-40. 2000 have some biogeographic generality. eas of Fennoscandia (see former discussion). Tenow & Nilssen (1990) demonstrated that the There are two features which may partially ex­ differences between the winter climate ofthe fjord plain the isolations ofthe beetles: 1) Continental districts of Finnmark and the areas further south parts ofTroms and the fjord districts ofFinnmark and east may be of great importance for insects have higher ambient summer temperatures than hibernating above the snow cover. Several soil­ the coastal districts and appreciably higher ambi­ dwelling terrestrial snail and slug species are also ent winter temperatures than northern Sweden and absent in this northfennoscandian «cold center». Finland (Table 2). Only in the coastal districts of Such species are e. g. Columella aspera Walden, the GulfofBothnia south ofabout 65° N are win­ Clausilia bidentata (Str0m), Limax marginatus ter as well as summer temperatures higher than in (Muller), 1. tenellus (Muller) and Arianta arbu­ the current areas in northern Norway (Table 2, Fig­ storum (L.) (Kerney & Cameron 1979, Andersen ure 6, see also Dahl 1998). Furthermore, the me­ 1982, Andersen & Halvorsen 1984). 2) North Nor­ teorological stations at 0verbygd and Bardufoss way has a high topographic reliefwhereas the ter­ are situated in the bottom ofvalleys. Winter tem­ rain becomes successively flatter from the Nor­ peratures are higher on the side of valleys than in way-Sweden border to the Gulf of Bothnia. In the bottom (see 2) below). Since the beetles of Dividalen E. lundbladi and C. pineti were found concern are saproxylic and live on or in stubs or on SW-facing sides of the valley. S and SW-fac­ standing or lying trees, it is likely that they would ing slopes have favourable microclimates with a be frequently exposed to temperatures close to the higher minimum and daytime temperature than ambient temperature in the most continental ar-

Table 2. Climatological data from meteorological stations in northernmost Norway and Sweden Data are from the period 1961-1990 and are taken from Aune (1993), F0rland (1993) Alexandersson' et al (1996). Tc: mean temperature in coldest month; Tw: mean temperature in July. ' .

Area Station Tc Tw Precipitation (mm) °C °C May-August Troms, coast Torsvag -1.1 10.8 206 Sommamy, Senja -1.9 11.9 245 Troms0 -4.4 11.8 266 Troms, inland Dividalen -9.4 12.8 147 Bardufoss -10.4 13.0 182 0verbygd -10.2 13.2 207 Finmark, fjords Alta -9.0 13.5 157 Banak -10.0 12.7 164 Finmark, inland Kautokeino -16.1 12.4 185 Karasjok -17.1 13.1 194 Pasvik -15.4 13.7 191 Sweden, northernmost part Abisko -11.7 11.3 134 Karesuando -16.0 12.8 208 Kiruna -13.8 12.1 240 Kvikkjok -15.2 12.6 262 Jokkmokk -16.6 14.2 235 Pajala -14.8 14.1 225 Coast of Gulf of Bothnia Haparanda -12.1 15.4 187 Lulea -11.5 15.5 185 Pitea -11.0 16.0 190 Umea -9.1 15.2 218

37 Andersen et al.: Saproxylic beetles (Coleoptera) from northern Norway

N-facing slopes and flat areas, e. g. bottom of and/or continental parts of Troms may be ex­ valleys. The former may therefore support a ther­ plained by these areas having comparatively fa­ mophilic fauna and flora (Krogerus 1960, Geiger vourable summer climate in combination with 1965, Munn 1966, Hulten 1971, Andersen & only moderately low winter temperatures. Hanssen 1989, Voisin 1990). In addition, lowpre­ cipitation in summer (May-August) in the current Acknowledgement. We are grateful to O. Hanssen and areas (Table 2) may also contribute to a favour­ F. 0degaard for verifying the identification ofEnicmus able rnicroclimate since sparse rainfall means less lundbladi and Corticaria pineti and for providing us wetting and, consequently, reduced cooling ofthe with infonnation about the distribution of beetles in substrate. Norway. We are also indebted to H. Rinden who, just before he tragically passed away, submitted a list of Thus, in continental parts ofTroms and the fjords beetles he had collected in Alta in 1998. S. Ligaard and ofFinnmark, comparatively favourable microcli­ T. Kvarnme likewise provided us with data about the mate in summer and moderately low winter tem­ distribution of beetles in Norway, and H. Silfverberg peratures may constitute an unique combination about the distribution of the beetles in Finland. A. within an area which otherwise either has low Granmo is acknowledged for infonnation about the dis­ tribution and ecology ofHypoxylon mammatum and R. summer temperatures or very low winter tempera­ Barret for correcting the English. tures.

What has been proposed here may partly be tested References experimentally. The cold hardiness (e. g. super­ Alexandersson, H., Karlstr0m, C. & Larsson-McCann, cooling point) of H. vilis, S. boleti, E. lundbladi S. 1996. Temperature and precipitation in Sweden and M. linearis should be tested with that ofmore 1961-90.87 pp. SMHI. Meteorologi 81,1991. or less close relatives which occur in the most con­ Aim, T. & Birks, H.H. 1991. Late Weichselian flora tinental parts of Fennoscandia, for instance and vegetation of And0ya, Northern Norway ­ Hapalarea linearis (Zetterstedt), Scaphisoma macrofossil (seed and fruit) evidence from Nedre subalpinum Reitter and Enicmus fungicola JErasvatn. Nord. 1. Bot. 11,465-476. Andersen, 1. 1982. New data on terrestrial gastropods Thomson. Furthermore, studies ofcold hardiness ofnorthem Nordland and Troms counties, N. Nor­ and further investigations of host selection of 0. way. Fauna norv. Ser. A 3, 37-40. festivus should be made. A comparable species Andersen, 1. & Halvorsen, O. 1984. Species composi­ might be 0. alni (Gyllenhal) which is distributed tion, abundance, habitat requirements and regional throughout Fennoscandia. More studies of most distribution of terrestrial gastropods in arctic Nor­ of the species in the field are needed. way. Polar BioI. 3, 45-53. Andersen,1. & Hansen, O. 1989. Billefaunaen i Gud­ CONCLUSION brandsdalen. Insekt-Nytt 14, 15-23. Audisio, P. 1993. Coleoptera Nitidulidae-Kateretidae. Six saproxylic species of Troms and western Fauna d Italia 32. 971 pp. Calderini, Bologna. Finnmark have exceptional distributions in Fenno­ Aune, B. 1993. Temperaturnonnaler, nonnalperioden scandia. Three of them have their main distribu­ 1961-1990. 63pp. Det Norske Meteorologiske Insti­ tion in the western part of this area. Orthocis tutt, Oslo. Bily, S. & Mehl, O. 1989. Longhorn beetles (Coleo­ festivus may be absent in northern Sweden and ptera, Cerambycidae) of Fennoscandia and Den­ Finland because of scarcity/absence of the host, mark. Fauna ent. Scand. 22, 1-203. Stereum rugosum there. Hapalarea vilis, which Brattli,1. G., Andersen, 1. & Nilssen, A. C. 1998. Pri­ has an atlantic distribution, may avoid continen­ mary attraction and host selection in deciduous and tal areas due to susceptibility to low winter tem­ conifer living Coleoptera: Scolytidae, Curculio­ peratures. The distribution pattern of H. ioptera nidae, Cerambycidae and Lymexylidae. 1. AppI. is enigmatic. The isolated northern occurrences EntomoI. 122, 345-352. of Scaphisoma boleti, Enicmus lundbladi and Coope, G. R. 1969. The contribution that the Coleoptera of Glacial Britain could have made to the subse­ Corticaria pineti in the fjords ofwestern Finnmark quent colonization of Scandinavia. Opusc. en!. 34,

38 Narw. J. Entama/. 41, 29-40. 2000

95-108. Koch, K. 1992. Die Kiifer Mitteleuropas. Okologie. 3. Dahl, E. 1950. Forelesninger over norsk plantegeografi. 389 pp. Goecke & Evers, Krefeld. 113 pp. Akademisk Trykningssentral, Oslo. Krogerus, R. 1960. Okologische Studien iiber nordische Dahl, E. 1992. Nunatakkteori. IV. Hvor fantes isfrie Moorarthropoden. Comment. BioI. 21,1-238. omnider og hva slags planter kunne leve pa dem? Lekander, H., Hejer-Petersen, B., Kangas, E. & Bakke, Blyttia 50, 23-35. A. 1977. The distribution ofbark beetles in the nor­ Dahl, E. 1998. The phytogeography of Northern Eu­ die countries. Acta Ent. Fenn. 32, 1-37. rope. 297 pp. Cambridge University Press, Cam­ Lid, J. 1985. Norsk-svensk-finsk flora. 837 pp. Det bridge. Norske Samlaget, Oslo. Eriksson, 1., Hjortstam, K. & Ryvarden, L. 1981. The Lindroth, C. H. 1949. Die fennoscandische Carabidae. Corticiaceae of North Europe. 6. Fungiflora. Uni­ Ill. Allgemeine Teil. Goteborgs K. Vetensk. - o. versitetsforiagets trykningssentral, Oslo. Vitterh. Samh. Handl. Ser. H 4, 1- 911. Erikson, 1., Hjortstam, K. & Ryvarden, L. 1984. The Lindroth, C. H. 1960. Catalogus Coleopterorum Corticiaceae of North Europe. 7. Fungiflora. Uni­ Fennoscandiae et Daniae. 476 pp. Entomologiska versitetsforlagets trykningssentral, Oslo. Sallskapet, Lund. Fossli, T.-E. & Andersen, 1. 1998. Host preference of Lucht, W. H. 1987. Die Kafer Mitteleuropas. Katalog. Cisidae (Coleoptera) on treeinhabiting fungi in 342 pp. Goecke & Evers, Krefeld. northern Norway. Entomol. Fennica 9, 65-78. Lundberg, S. 1964. Hidrag till Kannedom om Forland, E. 1. 1993. Nedbornormaler. Normalperioden skalbaggsfaunan pa Gotska Sand6n. Ent. Tidskr. 85, 1961-90. Rapport nr. 39/93 Klima. 63 pp. Det 45-48. Norske Meteorologiske Institutt, Oslo. Lundberg, S. 1969. Hidrag til kannedom om svenska Geiger, R. 1965. The climate near the ground. 611 pp. skalbaggar 12. Ent. Tidskr. 90, 217-229. Harvard University Press, Cambridge. Lundberg, S. 1995. Catalogus Coleopterorum Sueciae. Gj~revoll, O. 1992. Plantegeografi. 200 pp. Tapir, Naturhistoriska Riksmuseet & Entomologiska F6re­ Trondheim. ningen i Stockholm, Stockholm. Hansen, V. 1964. Fortegnelse over Danmarks biller Lysholm, H. 1937. Coleopterfaunaen iTrondelag. Norsk (Coleoptera). Ent. Meddr. 33, 1-506. ent. Tidsskr. 4, 145-182. Hansen, V. 1968. Biller 25. Atselbiller, stumpbiller m. Munn, R. E. 1966. Descriptive micrometeorology. 245 m. Danm. Fauna 77, 1-353. pp. Academic Press, New York. Haugen, L. 1999. En sammenligning av sprednings­ 0degaard, F. 1994. Kuldetilpasninger hos norske arter evnen hos ulike grupper av saproxyliske biller. 30 av trebukk- slekten Rhagium (Fabr.) (Coleoptera, pp. Cand. scient. thesis, University of Tromso, Cerambycidae). Er kuldetilpasninger en begrensen­ Tromso. de faktor for artenes utbredelse? 44 pp. Cand. sci­ Hulten, E. 1971. Atlas over vaxternas utbredning i Nor­ ent. thesis, Norwegian University of Science and den. Fanerogamer och ormbunkvaxter. 531 pp. Technology, Trondheim. Generalstabens LitografiskaAnstalts Forlag, Stock­ 0kland, K. A. 1981. Inndeling av Norge til bruk ved holm. biogeografiske oppgaver - et revidert Strand-sys­ Huse, 0. 1997. Kan mengden ynglemateriale i et skogs­ tem. Fauna (Oslo) 34,167-178. ornrade gi informasjon om artsrikdom og individtall Olofsson, D. 1996. Tickor i Sverige. 127 pp. WWF. av obligatorisk saproxyliske biller? 46 pp. Cand. SCA Skog AB, Sundsvall, Sverige. scient. thesis, University of Tromso, Tromso. Palm, T. 1959. Die Holz- und Rindenkafer der siid- und Iversen, M. 1997. Vednedbrytende sopp pa graor (Alnus mittelschwedischen Laubbaume. Opusc. ent. Suppl. incana (L.) Moench) i Reisadalen nasjonalpark­ 16,1-374. artssammensetning, artsrikdom og okologi. 58 pp. Ryman, S. & Holmasen, I. 1992. Svampar. En falthand­ Cand. scientthesis, University ofTromso, Tromso. bok. 718 pp. Interpublishing, Stockholm. Johnson, C. 1967. Coleoptera in southern Nordland. Ryvarden, L. 1971. The genera Stereum (s. lato) and Norsk ent. Tidsskr. 14,70-82. Hymenochaete in Norway. Norw. 1. Bot. 18, 97­ Kerney, M. P. & Cameron, R. A. D. 1979. A field guide 108. to the land snails of Britain and North-West Eu­ Ryvarden, L. 1976. The Polyporaceae ofNorth Europe. rope. 288 pp. Collins, London. 1. Albatrellus- Incrustoporia. 1-214. Fungiflora, Koch, K. 1989a. Die Kafer Mitteleuropas. Okologie. Universitetsforlagets trykningssentral, Oslo. I. 440 pp. Goecke & Evers, Krefeld. Solheim, H. 1979. Vedboende sopp (Aphyllophorales, Koch, K. 1989b. Die Kiifer Mitteleuropas. Okologie. Homobasidiomycetes) i to omrader i Engerdal, 2. 382 pp. Goecke & Evers, Krefeld. Hedmark (Gutulia nasjonalpark og Hovden) og i

39 Andersen et al.: Saproxylic beetles (Coleoptera) from northern Norway

0vre Dividalen nasjonalpark, Troms. 81 pp. The­ sis, University of Oslo. Strand, A. 1946. Nord-Norges Coleoptera. Troms" Mus. Arsh. 67 (1944),1-629. Strand, A. 1953. Coleoptera fra Nordreisa. Norsk ent. Tidsskr. 4, 63- 70. Strand, A. 1961. En overraskende forekomst av Antho­ bium sorbi Gyll. i Nord-Norge (Col., Staphylini­ dae). Norsk ent. Tidsskr. 11,235- 239. Strand, A. 1970. Additions and corrections to the nor­ wegian part of Catalogus Coleopterorum Fenno­ scandiae et Daniae. Norsk ent. Tidsskr. 17, 125­ 145. Strand, A. 1977. Additions and corrections to the nor­ wegian part of Catalogus Coleopterorum Fenno­ scandiae et Daniae. Second series. Norw. J. Ent. 24, 159-165. Strid, A. 1975. Wood-inhabiting fungi of alder forests in North-Central Scandinavia. I. Aphyllophorales (Basidiomycetes). Taxonomy, ecology and distri­ bution. Wahlenbergia 1, 1-237. Tenow, O. & Nilssen, A. 1990. Egg cold hardiness and topoclimatic limitations to outbreaks of Epirrita autumnata in Northern Fennoscandia. J. Appl. Ecol. 27, 723-734. Vik, A. 1995. Lathridiidae (muggbiller). Norske Insekt­ tabeller 15. 15 pp. Norsk Entomologisk Forening, Oslo. Voisin, J.-F. 1990. Chorthippus biguttulus (Linnaeus 1758) funnet i Gudbrandsdalen (On) (Orthoptera: Acrididae). Fauna norv. Ser. B. 37, 47. Vorren, T. 0., Vorren, K.-D., Aim, T., Gulliksen, S. & L"vlie, R. 1988. The last deglaciation (20.000­ 11.000 B. P.) on Andeya, northern Norway. Boreas 17,41-77.

Received 15 September 1999, accepted 30 November 1999

40 Norw. J. Entomol. 41, 41-63. 2000

A review of the terrestrial and freshwater invertebrate fauna of the High Arctic archipelago of Svalbard

Stephen J. Coulson

Coulson, S.l 2000. A review of the terrestrial and freshwater invertebrate fauna of the High Arctic archipelago of Svalbard. Norw. J. Entomol. 47,41-63.

The terrestrial and freshwater invertebrates of Svalbard (Spitsbergen) are reviewed. In order to pro­ vide as complete an overview ofthe invertebrate fauna as possible this article is not restricted solely to the Insecta, but includes the phyla Rhizopoda, Rotifera, Tardigrada, Nematoda, Annelida, Crusta­ cea and also the classes Collembola and Arachnida. Population densities and dynamics, as well as physiological responses to the high arctic environment, are discussed. Species diversity is poor on Svalbard by comparison with other arctic regions. It is suggested that this is due to a combination of the extreme isolation of the archipelago, the high Arctic climate and the ecologically young age of the ecosystem. The distribution of the invertebrate species throughout the islands is dependent on factors such as microclimate, vegetation, bed rock and water quality. The majority of the inverte­ brate species have wide circumpolar distributions and there are few endemics. The four principle methods of dispersal to islands; i) wind currents (and frontal systems), ii) oceanic currents (drift wood etc.), iii) animal vectors and iv) human introductions, are discussed with relation to the current Svalbard fauna. The inhabitation of glacial refugia on Svalbard during the last glaciation is consid­ ered.

Key words: invertebrate, Svalbard, Spitsbergen, dispersal, Arctic.

Department ofBiology, Division ofZoology, University ofOslo, PO Box 1050 BUndem, N-0316 Oslo, Norway. Current address: Division ofBiological and Earth Sciences, Liverpool John Moores University, Byrom St., Liverpool, L3 3AF, UK.

INTRODUCTION press) presents the most complete overview ofthe invertebrate species and ecology since S",mrne Svalbard is a High Arctic archipelago situated (1979) and highlights key aspects of this fauna. between 74°N to 81 oN and 10 0 E to 35°E, The is­ 2 lands have an area ofapproximately 63,000 km , Such an overview is required due to the rapidly 60% of which is permanently covered by ice or increasing number of known species, the abun­ snow (HisdaI1985, Mangerud & Svendsen 1992). dance of synonyms in the literature and the rising The climate ofSvalbard is unusually mild despite number ofinvertebrate ecological and ecophysio­ the high latitude location due to the warming in­ logical studies being undertaken on Svalbard (for fluences of the Atlantic current and southerly air example Leinaas & Arnbrose 1992, Strathdee et streams (Hisdal 1985, Mangerud & Svendsen al. 1993a, Hertzberg et al. 1994, Ho1mstrup & 1992, Fogg 1998, Danks 1999). This, combined S",mme 1998, Hoba::k & Weider 1999, S",mme & with the geographic isolation of the islands, has Birkemoe 1999, Coulson et aI., in press). Some of resulted in a faunal assemblage that is quite dis­ the first collections of insect from the region of tinct from other arctic regions. This article which Svalbard were taken in 1836 (Boheman 1865 in is based on the checklist ofCoulson & Refseth (in S",mme (1993)) and by 1930 some 300 species of

4 1 eoulsOn: Invertebrate fauna of the High Arctic archipelago of Svalbard

30 0­ 20 'L­ 10 ~ ~ .3 o t- - -- 7 , ~ Q) -10 Cl. E -20 .---.--- Q) Figure 1. Mean 30 year monthly air I- -30 temperatures (1957-76) at Longyear­ byen airport (e), maximum monthly air -40 temperatures (.) and minimum monthly -50 I i i air temperatures (A) during the same 1 Jan. 1 Mar. 1 May 1 July 1 Sept. 1 Nov period. (Data from F0rland et al. 1997b).

terrestrial or freshwater invertebrate known from been recorded, for example 21.3°C at Svalbard Svalbard (Thor 1930). This number has since airport (Longyearbyen) in July 1979 (F0rland et grown to 1040 (Coulson & Refseth, in press). This al. 1997b). The upper soil layers are typically is however, far from being a complete description thawed for only three months, mid-June to mid­ of the actual total number of invertebrate species September, (Coulson et al. 1995) with a total tem­ present due to i) the majority of collections have perature sum of between 200 to 400 degree days been made at, or in the vicinity of, only three lo­ by late August (Coulson et al. 1993). Soil tem­ cations on the west coast of Spitsbergen (Long­ peratures are partially dependent on air tempera­ yearbyen, Ny-Alesund and Homsund; but see for ture but also on insolation (Coulson et al. 1993). example Summerhayes & Elton 1928, Dastych Temperatures in the upper 3 cm are often above 1985, Fjellberg 1997, Hauge & S0mme 1997 for 10°C (Coulson et al. 1993) while temperatures exceptions), ii) the likelihood of a bias towards above 20°C in the soil surface/vegetation inter­ those species easily identified to the detriment of face are not uncommon and can peak at over 30°C those species difficult to collect and/or identify, (Coulson et al. 1993, Hodkinson et al. 1996b, iii) concentration on those taxa that have caught Coulson, Leinaas & lms, unpub. data). the interest ofarctic taxonomists, and iv) misidenti­ Mean winter air temperatures on the west coast of fications, confusion over synonyms and inconsist­ Svalbard are seldom below -20°C (Figure 1) and ency in reference works. soil temperatures are typically between -5 and ­ This article seeks to provide a general overview 20°C (Coulson et al. 1995, Coulson et aI., in press). of the invertebrate species assemblage found on This is partly due to the warm West Spitsbergen Svalbard, discuss unusual aspects ofthis fauna and current (>3 0c), a northwards extension of the present possible explanations for species presence North Atlantic Drift, flowing up the west coast and absence. (Fogg 1998) (Figure 2), but in addition, the pas­ sage of low pressure weather systems over Sval­ Physical features of Svalbard bard also transport large volumes ofcomparatively warm air from more southerly latitudes (Hisdal Climate 1985). By comparison, mean winter air tempera­ The summer period is short and only four months tures in Greenland are below -30°C for several have positive mean air temperatures (Figure 1). months each year (Fogg 1998). Minimum winter Mean normal summer air temperatures in the soil temperatures on Svalbard are dependent on warmest months (July and August) are in the or­ the depth ofthe snow cover but typically vary from der of 4 to 5°C (Fflrland et al. 1997b) but on rare -30°C (no snow cover) to -5°C (snow cover) occasions much higher peak temperatures have (Coulson et al. 1995, in press). The period of the

42 Norw. J. Enlomol. 41, 41-63. 2000

Sju0yane 0° West Spitsbergen r wsc \

Figure 2. Map of Svalbard with the ocean currents marked and posi­ tions of the three principal study areas, Longyearbyen (LYB), Ny­ ESC Alesund (N-A) and Hornsund indi­ cated. WSC = West Spitsbergen current, ESC = East Spitsbergen current.

midnight sun is from mid-April to late August Vegetation, soils and water quality (Hisdal1985). The east coast ofSvalbard is colder Although 2885 species ofplants, algae, fungi and by virtue of a cold southerly flowing current that cyanobacteria are known from Svalbard (Elvebakk originates in the Russian arctic as the Transpolar & Prestrud 1996) much of the coastal region can Drift (Fogg 1998). Mean monthly summer air tem­ be described as either polar semi-desert or tundra peratures along the east coast are between 1.5 to heath type vegetation assemblages. The polar 2.5 deg. C lower than on the west coast (Hisdal semi-desert is characterised by large areas ofbare 1998). soil and or stones with a thin surface covering of Precipitation in the arctic is generally low (Ferland cyanobacteria and lichens with vascular plants et al. 1997b). On the west coast of Svalbard an­ such as Dryas octopetala. Organic soils are thin, Dual precipitation is between 190 mm and 480 mm often less than 1 cm thick. Such habitats are typi­ (at Svalbard airport and Isfjord Radio respec­ cal of well drained glacial moraines (Coulson et tively). Note the large local variation in precipita­ a1. 1993). In contrast, the tundra heaths are often tion. Isfjord Radio is only 50 km distant from moister with a better developed organic soil up to Svalbard airport. Large differences can also occur 10 cm in depth. Vascular plants such as Cassiope on a more local scale. Ferland (et a1. 1997a) meas­ tetragona are emergent through a layer of moss ured 60% greater precipitation on the Austre Bmg­ (often Racomitrium spp.) (Coulson et a1. 1993). gerbreen (East Bmgger glacier) than at the settle­ In low lying poorly drained areas there is often a ment of Ny-Alesund only 2.5 km distant. Such thick layer ofcold wet moss. For instance, beneath differences have implications for soil moisture the Stuphallet bird cliffs the moss layer is ca. 10 contents, temperature and vegetation type with cm thick (Cooper 1996). consequent effects on the distribution of the in­ The majority of the mineral soils are gravels for­ ,.r vertebrate fauna. 43 eoulson: Invertebrate fauna of the High Arctic archipelago of Svalbard med from eroded glacial moraine during the re­ 1994). By comparison with more temperate re­ cent Quaternary. Sand and clays are only found gions this is a low total and is comprised largely in the river valleys (Hjelle 1993). The active layer of cosmopolitan species (Beyens et al 1986a, b, is thin, often about 1.5 m in depth, over a perma­ Beyens & Chardez 1995, Beyens & Chardez frost which extends to a depth of between 100 to 1997), with the genus Centropyxis dominating 460 m (Hjelle 1993). Periglaciallandforms such (Beyens & Chardez 1995). Note however the grea­ as solifluction lobes, rock glaciers and sorted ter species richness ofSvalbard compared to some ground are common. Antarctic island groups (Balik 1994). Chardez & Beyens (1987) comment on both the low species INVERTEBRATE TAXA number and low diversity indices ofsamples from Edgeoya. This they attribute to the extreme high Phylum Rhizopoda arctic conditions and the fact that the samples came The testate amoebae are represented by 190 spe­ from a wet moss habitat which represents the most cies on Svalbard (Table I) inhabiting aquatic, moss favourable habitat available. and soil habitats (Beyens et al 1986a,b, Balik

Table 1. Number of freshwater and terrestrial invertebrate species from Svalbard and including the number of insect families from Svalbard and Greenland. ('dubious record, J0rgensen and Eie (1993), "total includes the dubious ephemeropteran). Total number of Phyla Class Order species on Svalbard Rhizopoda 190 Actinopoda 3 Ciliophora 6 Apicomplexa 1 Rotifera 153 Gastrotricha 1 Nematoda 108 Platyhelminthes 8 Annelida 34 Tardigrada 83 Chelicerata Arachnida Acari 116 Araneae 18 Mandibulata Collembola 59 Insecta Anoplura 2 Hemiptera 4 Mallophaga 37 Dictuoptera 0 Dermaptera 0 Coleoptera 19 Diptera 128 Lepidoptera 11 Hymenoptera 26 Siphonaptera 2 Trichoptera 1 Ephemeroptera "1 Crustacea Branchiopoda 9 Copepoda 10 Malacostraca 9 Ostracoda 2 Total **1040

44 Norw. J. Entomol. 47, 41-63. 2000

Phylum Rotifera tribution (Dastych 1985). For example, Ramaz­ A total 153 species ofRotifera have been recorded zottius cataphractus (Hypsibius cataphractus) is from Svalbard (Table 1). Studying Keratella only found at altitudes below 100 m while Hebe­ quadrata Amren, (1964) found that the animals suncus conjungens (Diphascon conjugens) is re­ from Svalbard had a higher specific metabolism stricted to areas above 1100 m. Dastych (1985) than temperate strains. The metabolism of two comments on the similarities in the distribution common rotifers from Svalbard, Macrotrachela with species assemblages from the Tatra moun­ musculosa and Trichotria truncata, have a low tains. Each cryoconite hole can be considered an temperature dependency over a range oftempera­ individual ecosystem and can possess distinct fau­ tures typical ofSvalbard water temperatures (Opa­ nas ofRotifera and Tardigrada (such as Diaphas­ linsky & Klekowski 1989). Amren (1964) also con recamieri and Isohypsibius granulifera) which observed morphological differences in the genera­ often occur in large numbers (de Smet & van tions of K. quadrata. The animals overwinter as Rompu 1994). There are few records oftardigrade dormant eggs and the first generation has a dis­ density but Wiithrich (1992) observed densities tinctive morphology. Eggs are produced in a set below 5,000 m- 2 in oligotrophic tundra. sequence beginning with amictic eggs resulting in a rapid population increase, but with mictic eggs Phylum Nematoda later in the season. Due to high overwintering egg There have been no quantitative ecological or phy­ survival, initial densities are ca. 40 individuals per siological studies ofthe nematodes from Svalbard. litre (Amren 1964). Cryoconite holes form on the However, Loof (1971) identified 74 species of surface of glaciers due to localised melting fol­ free-living or plant parasitic nematode from an lowing solar warming of dark detritus. Large extensive sampling program on West Spitsbergen. populations of Rotifera often occur in such holes, Including the vertebrate parasites the total nema­ for example Macrotrachela insolata and Philodina tode fauna numbers 108 species (Table 1). Loof acuticornis odiosa, which are predated by larger (1971) highlights the lack of diversity between Rotifera (de Smet & van Rompu 1994). de Smet habitats and the absence of many families and (1993) commented on how well the Rotifera are genera on Svalbard that are widespread in Europe. adapted to maximise dispersal. A high reproduc­ For example, the family Rhabditidae and genus tive rate, parthenogenicity and the production of Rotylenchus. For reasons that are unclear, the desiccation resistant eggs all favouring dispersal nematode Geocenamus microdorus is restricted to by both wind and birds. the small islands of Eskjeret and Juttahl (small is­ lands opposite Ossiansarsfjellet in Kongsfjord) Phylum Tardigrada where it often forms the dominant species (Loof The Tardigrada is a widely distributed group on 1971). The nematode fauna of Svalbard appears Svalbard with 99 species recorded (Table 1). Clo­ to have close affinities to that ofnorthern Canada sely situated and similar habitats may reveal dis­ (Loof 1971). similar species assemblages (Dastych 1985, Mauc­ ci 1996, van Rompu & de Smet 1996). Dastych Cestode and nematode parasites (1985) demonstrated that the species composition The parasitic Cestoda and Nematoda are discussed at a locality is dependent on a combination ofbed­ as one group due to their mode of life although rock type and habitat altitude. Tardigrade species they do not belong to one taxon. can be classified into five groups based on bed rock preferences from eucalciphilous (e.g. Macro­ In addition to man, the terrestrial mammal fauna biotus willardi) through mesocalciphilous (e.g. of Svalbard is restricted to five species (Mehlum Hypsibius oberhaeuseri) to acalciphilous (e.g. H. 1990); reindeer (Rangifer tarandus platyrhyn­ dujardini). Furthermore, numbers ofboth individu­ chus), arctic fox (Alopex lagopus), polar bear (Ur­ als and species decrease on an altitudinal gradi­ sus maritimus, more strictly a marine mammal), ent, many species having a distinct altitudinal dis­ the vole (Microtus rossiameridinalis) and dogs

45 COl/lson: Invertebrate fal/na of the High Arctic archipelago of Svalbard

(Canis domesticus, largely Greenland sled dogs). parasites on Svalbard during autumn and late win­ Several other species, the muskox (Ovibos mos­ ter has been recently demonstrated and is the first chatus), arctic hare (Lepus arcticus), mountain such observation during winter in High Arctic hare (L. timidus), Norwegian rat (Rattus nor­ environments (Halvorsen et al. 1999). It should vegicus), mouse (Mus musculus domesticus) and be noted that many ofthe trichostrongyloid nema­ one fish, the trout (Salmo trutta), have all been todes are now considered to be dimorphic or tri­ introduced during the last century but have failed morphic species with major and minor morphoty­ to establish viable populations (Gjertz & Lene pes (Halvorsen & Bye 1999). Infection rates of 1998). While many of these species have inverte­ Trichinella spp. can be high, infection rates in the brate parasites the total number of parasite spe­ polar bear vary between 23 and 58% and from 3 cies is low, one reason being the lack of suitable to 67% in the fox. However no infection was re­ intermediate hosts. For example, the lack of tre­ ported in samples from 252 ringed seals (Phoca matodes and metastrongyloid nematode species hispida) and 84 bearded seals (Erignathus bar­ among non-migrating host species can be explai­ batus) (Larsen& Kjos-Hanssen 1983). Extensive ned by the lack of gastropods on Svalbard (Hal­ freeze resistance has been demonstrated in Trichi­ vorsen & Bye 1999). nella isolates with live specimens being obtained from polar bear muscle after either 38 months at ­ Cestoda 18°C or 7 weeks at -70°C (Kjos-Hanssen 1984). In contrast, Trichinella from more temperate re­ Eight cestodes (Platyhelminthes: Cestoda) para­ sitising the arctic charr (Salvelinius alpinus) or the gions is far more cold susceptible. Pork is rou­ tinely frozen at -15°C for 20 days to kill Trichinella reindeer (R. tarandus platyrhynchus) are known cysts and prevent transmission of the nematode from Svalbard; five from the arctic charr and two (Kjos-Hanssen 1983). Nematode infection of the from the reindeer. Furthermore, Echinococcus glaucous gull (L. hyperboreous) has a positive re­ multilocularis (Cestoda: Taeniidae) has been re­ lationship with tissue polychlorinated biphenyl cently observed amongst the voles (Microtus (PCB) concentrations (Sagerup et al. 1998). Sup­ rossiameridinalis) at the derelict Russian mining pression of the immune system by the PCB load settlement of Grumant (lsfjord) (R.A. Ims & H. is thought to increase susceptibility to parasite in­ Henttonen, pers. comm.). The definitive host of fection. this parasite is the fox, the vole forming the inter­ mediate host. On occasion man can form the inter­ One additional parasitic nematode, Philonema mediate host and in such instances infection with oncorhynchi, occurs in conjunction with the arc­ the cyst usually results in alveolar hydatid disease tic charr (Kennedy 1978, Sobecka & Piasecki and gradual destruction of the liver. 1993). The free-living and plant parasitic nema­ todes from a wide ranging sampling program cov­ Nematoda ering most ofWest Spitsbergen have been detailed A total of 10 trichostrongyloid nematodes (Nema­ by Loof (1971). toda: Strongylida) have been recorded, nine from the reindeer and muskox (note that the muskox is Phylum Annelida no longer present on Svalbard) (Alendal & Helle There are no lumbricid worms on Svalbard. One 1983, Halvorsen & Bye 1999), and one from the immature form has been reported (Nurminen glaucous gull (Larus hyperboreus) (Sagerup et al. 1965) but this must be considered a dubious record 1998). Trichinella spiralis and T. nativa (Nema­ considering the large number ofsoil samples since toda: Adenophorea) are prevalent in both the po­ taken and no further records emerging. However, lar bear (Ursus maritimus) (Kjos-Hanssen 1984) 34 species ofenchytraeid are known from Svalbard and the arctic fox (Alopex lagopus) (Prestrud et (D6zsa-Farkas 1999). The species present have a al. 1993). wide circumpolar distribution, but there is con­ siderable nomenclature confusion in the literature The continued transmission ofnematode reindeer

46 Norw. J. Enlomo/. 41, 41-63. 2000 on the Enchytraeidae largely due to the identifi­ known about the life history ofmany arctic mites cation problems. D6zsa-Farkas (1999) has recently but it seems likely that, as with some Antarctic published a review that resolves these confusions. species (Convey 1996), many species from Sval­ bard require several years to attain sexual maturi­ There are few reports dealing with the ecology ty. Mites have a greater tolerance to climatic stres­ and physiology of the enchytraeids (Birkemoe ses such as high soil temperatures and drought than 1995, S0mme & Birkemoe 1997, Birkemoe et al. Collembola (Coulson et al. 1996, Hodkinson et submitted). The life cycle typically takes two years al. 1996b), but the slow generation time will de­ to complete on Svalbard, for example Henlea glan­ lay their response to climate change. Very little is dulifera. Other species such as Henlea perpusilla known concerning the ecology and ecophysiology have the ability to mature in one year. In less ex­ of the Acari fauna of Svalbard. treme environments H. perpusilla can complete two generations a year (Dash & Cragg 1972). The cocoon is an important overwintering stage and, Order Araneae although cocoons are produced throughout the The spiders are better known than the mites and year, cocoon hatch is synchronised in the spring 18 species from three families are represented on by low winter temperatures breaking a form of Svalbard, the majority from the family Linyphiidae quiescence (Birkemoe et al. submitted). Birkemoe and only single species from each ofthe Gnapho­ et al. (submitted) also failed to observe reproduc­ sidae and Hahniidae. Two species may be human tion by fragmentation amongst the five species of introductions, Tapinocyba insecta (Linyphiidae) enchytraeid studied. Overwintering enchytraeids and Hahnia helveola (Hahniidae) (Tambs-Lyche are freezing susceptible (S0mme & Birkemoe 1967). The remaining species all have circumpolar 1997). However, the enchytraeids can cold accli­ distributions. While the total number of species mate and desiccate, loosing up to 73% ofthe body may be restricted, densities can be greater than in water content, and overwinter in this anhydrobiotic Finnish Lapland (Koponen 1980). Spiders are state (S0mme & Birkemoe 1997). common throughout Svalbard, one species (Collinsia spetsbergensis) even being recently col­ Phylum Chelicerata lected from the isolated islands of Sju0yane off 0 Order Acari the northern coast of Svalbard and lying at 80 N (Hauge & S0mme 1997) (Figure 2). Hagvar & There are 116 species of Acari on Svalbard (98 Hegstad (1969) point out that the wide habitat tol­ Acariformes and 18 Parasitiformes) (Table 1). The erance of spiders may be considered a pre-adap­ oribatid mites (Oribatida) are perhaps the best tation to the Svalbard environment where the land­ known group with 63 species. However, due to scape consists ofa mosaic ofdifferent habitats with synonyms and misidentifications, the actual num­ small areal extents (Leinaas 1999a). ber ofspecies present on Sva1bard is possibly only half this number (T. Solh0Y, pers. comm.). Dia­ Phylum Mandibulata pterobates notatus appears to be one of the most widespread species (Neidbala 1971). The classes Diplura or Protura are not found on Svalbard but there are 59 species of Collembola. Mite densities are in general low when compared to the Collembola (Table 2). For example, while Class Collembola Bengtson et al. (1974) recorded mite densities of 2 Of the Svalbard Collembola, 58 species have a between 16,000 and 248,000 mites m- , Wiithrich holarctic or circumpolar distribution. For exam­ (1992) found densities below 2,000 m-2• Prior to moulting mites enter a quiescent stationary stage. ple, Megalothorax minimus has a largely global It is unknown how this may effect density estima­ distribution including the Antarctic. Only Mesa­ tes that are based on samples from MacFadyen phorura jirii has a distribution restricted to Eu­ type extractors in which the soil microarthropods rope and Bj0rn0ya (Janssens 2000). In the absence are expelled and are required to be active. Little is oflumbricid worms on Svalbard the remaining soil

47 Coulson: Invertebrate fauna of the High Arctic archipelago of Svalbard

3 Z Table 2. Density (10 m' ) of some invertebrates on Svalbard. Note: Such density measurements are dependent on the sampling date and sampling and extraction methodologies. a Cryptostigmata only; b Collembola and Acari combined.

Habitat type Collem- Acari Nema- Enchytrae- Araneae Diptera Coleoptera Source bola toda idae larvae lichen tundra 9 16 0.07 0.06 Bengtson et al. 1974 Moss-lichen tundra 27 19 0.05 0.04 Bengtson et al. 1974 Wet moss tundra (birdcliffs) 243 22 2.5 Bengtson et al. 1974 Grassland 267 248 1.0 0.01 0.3 Bengtson et al. 1974 Tundra heath 10 Birkemoe et al. subm. Polygonal tundra 5 - 35 5 - 30 1.3 - 5.5 0.05 0.02 Byzova et al. 1995 Wet moss tundra 270 - 432 38 - 90 4.6 - 20 0-0.2 Byzova et al. 1995 lichenous tundra 432 - 104 31 -52 6.4 - 24 0.06 0.004 Byzova et al. 1995 Wet moss tundra. birdcliffs 266 - 592 39 - 81 0.2 - 7.6 0.06 0.008 0.06 Byzova et al. 1995 Polar semi-desert 30 - 83 4 - 14 Coulson et al. 1996 Tundra heath 14 - 40 5 - 11 Coulson et al. 1996 Carex tussocks 240 - 290 Hertzberg et al. 1994 Cyanobacterial ground 9 - 30 Hertzberg et al. 1994 Unknown 270 66 2.3 5.5 0.02 0.4 0.01 Klekowski & Opalinsky 1990 Tetragona-Dryadetum (inner fjord) 4 Sendstad 1976 Tetragona-Dryadetum (outer fjord) 19 Sendstad 1976 Polari-Dryadetum 8 Sendstad 1976 Caricetum ursinae 18 Sendstad 1976 lichen tundra 27 32' 10 Sendstad & Sveum 1985 Wet tundra 24 37' Sendstad & Sveum 1985 Dryas tundra 20 37- 7 Sendstad & Sveum 1985 Moss tundra 112 53- 4 Sendstad & Sveum 1985 Beach 65 20- 2 Sendstad & Sveum 1985 Moss-lichen tundra 18 - 81 Sendstad & Sveum 1985 Moss near beach 246 Somme & Birkemoe 1999 Moss tundra 231 Somme & Birkemoe 1999 Bottom of birdcliffs 80 Somme & Birkemoe 1999 Under birdcliffs 65 Somme & Birkemoe 1999 Rock shelves 68 Somme & Birkemoe 1999 Birdcliffs 224 22.5 376 100 3 0.2 WOthrich 1989 Oligotrophic tundra 40.2 6.7 81.1 <0.3 <0.03 WOthrich 1989 Open ground 3.7 0.3 WOthrich 1992 Oxyriazone 1.9 3.3 WOthrich 1992 Saxifrage zone 1.3 1.8 WOthrich 1992 Salixzone 0.6 0.3 WOthrich 1992 Moss tundra 1.9 0.8 WOthrich 1992 Moss tundra B.5 b WOthrich 1991 Cassiope tundra 11.8b WOthrich 1991 Salixtundra 5.3b WOthrich 1991 Festuca tundra 10.8b WOthrich 1991

fauna takes on an increased importance in decom­ the purple sandpiper (Calidris maritima) (Hagvar position processes and nutrient recycling (Lavelle 1970, Leinaas & Ambrose 1992, 1999). et al. 1997). It is likely that the Collembola and The density ofCollembola (and species dominan­ mites have an important role in these processes ce) varies greatly between habitats (Table 2). but there is a dearth ofpolar studies. Collembola, Hertzberg et al. (1994) observed Collembola for example Archisotoma megalops, may also be densities of between 9,000 to 290,000 individu­ important as part ofthe diet ofthe breeding wader als m-2 from open cyanobacterial ground and

48 Norw. J. Entomol. 41, 41-63. 2000

Carex tussocks respectively while Somme & Hypogastrura tullbergi has a synchronised life Birkemoe (1999) found densities of Folsomia cycle (Birkemoe & Leinaas 1999), the phenology quadrioculata of 85,000 m-2 and F. bisetosa of of which is maintained by a winter diapause ter­ 188,800 m-2 from Krykkjefjellet (kittywake bird minated by low winter temperatures. This ensures cliffs in Kongsfjord). In exceptional areas, such egg hatch after the spring thaw, a period of high as wet Carex tundra, population densities can ap­ nutrient availability and avoids the hatchlings proach 800,000 individuals m-2 (pers. obs.). Al­ overwintering (Birkemoe & Leinaas 1999). though some taxa are rare on Svalbard, Bengtson Some species show a distinct diel periodicity. For et al. (1974) concluded that arthropod densities example, Isotoma anglicana exhibits distinct on Svalbard were little different from high alpine peaks in activity, being most active in either early regions in mainland Norway. As well as varying morning or late afternoon. In contrast, Ceratophy­ spatially, the density of Collembola varies tem­ sella longispina (Hypogastrura hirsuta) was ar­ porally (Birkemoe & Somme 1998, Coulson et rhythmic (Solem & Sendstad 1978). This response al. 1996, Somme & Birkemoe 1999, Webb et al. is complicated, some species requiring diel tempe­ 1998). For example, from being highly abundant rature differences ofgreater than 5 deg. C to main­ in the years 1992 and 1993 (ca. 100,000 individu­ 2 tain synchronisation (e.g. Sminthurides malm­ als m- ), the density ofF. quadrioculata collapsed greni). It is hypothesised that the rhythmicity may in 1993 only to recover and then fluctuate in 1995 have great importance in providing a temporal as and subsequent years (Leinaas 1999b). Such tem­ well as spatial partitioning of niches. This may poral variation in density is largely due to the ef­ enable the co-existance ofclosely related species fects of stochastic environmental events. For ex­ (Solem & Sendstad 1978). In this context, differ­ ample, the formation ofa 25 cm thick winter sur­ ing tolerances to environmental stresses may also face ice layer, an event not uncommon in this re­ result in two closely related species utilising dif­ gion of the Arctic, reduced the population of F. ferent niches, for example the greater desiccation quadrioculata and Hypogastrura tullbergi by up resistance ofF. quadrioculata compared to F. sexo­ to 50% (Coulson et aI., in press). Summer condi­ culata (Hertzberg & Leinaas 1998). tions can also influence population performance. A three year cloche study of the responses of the The physiology ofthe collembolan Megaphorura soil fauna to elevated soil temperatures resulted arctica (often referred to as Onychiurus arcticus in a decline in Collembola densities (Coulson et or Protophorura arctica) is among the best stud­ al. 1996). This was attributed to the reduced soil ied ofthe invertebrates from Svalbard. Winter soil moisture status associated with the warmed soil temperatures can decline to -30°C (Coulson et al. and the inability ofthe Collembola to tolerate such 1995) and such extremely low temperatures pose dry environments (Coulson et al. 1996, Hodkinson a potential problem to the successful overwinter­ et al. 1998). ing of the Collembola. M. arctica, common in moist areas with a high nitrogen input (for exam­ The life cycle ofthese high arctic Collembola ap­ ple, amongst the vegetation beneath bird cliffs), pears to be extended by comparison to tempera­ has been shown to utilise a form ofanhydrobiosis ture relatives. The life cycles of two species, F. to survive these temperatures. It is currently the quadrioculata and H. tullbergi have been de­ only known collembolan (Holmstrup & Somme scribed in detail (Birkemoe & Somme 1998). 1998, Worland et al. 1998) to use such a strategy. While the exact life cycle differs between habi­ The collembolan avoids freezing even at tempera­ tats, F. quadrioculata on Svalbard usually requires tures as low as -30°C by reducing body fresh­ two years to reach sexual maturity and juveniles weight water content by up to 90% (Holmstrup & are found throughout the year (Somme & Birke­ Somme 1998). The acclimatory responses of the moe 1999). This is in contrast to F. quadrioculata enzyme amylase from M arctica and a temper­ from warmer more southerly latitudes where de­ ate species, Protophorura armata (Onychuirus velopment may be fast enough to permit up to armatus), show distinct differences (Sustr & Block three generations per year (Gregoire-Wibo 1976).

49 Coulson: Invertebrate fauna of the High Arctic archipelago of Svalbard

1998). While the temperature optimum ofP arma­ sect species respectively (Kevan & Danks 1986b). ta remains at 40°C, the activity ofthe enzyme from In an article considering the ornithology of Bear M arctica is dependent on the thermal acclimation Island, Munch (1962) mentions Thysanura but this of the insect. The temperature optimum from is an isolated record and seems likely to be a warm acclimated individuals is 35°C while after misidentification. nine weeks ofcold acclimation (5°C) the tempera­ ture declined to 20°C (Sustr & Block 1998). This Order Dip/era species also has a remarkably high QlO of 7 be­ Of the Mandibulata, it is the Diptera which has tween 0 and 10°C (Block et al. 1994). the greatest number of species, 128 species from The colder island ofNordaustlandet is notable for 21 families. However, many of the families from its lack ofspecies diversity (Leinaas 1999a). Nev­ Svalbard are poorly known. The Chironomidae ertheless, some species, for example the col­ in particular require revision, Lindegaard (1997) lembolan F quadriocu/ata, are particularly abun­ providing the most recent and accurate checklist dant. The reduced number of other species may (K. Aagaard, pers. comm.). To date, 63 species of have resulted in lower interspecific competition Chironomidae have been recorded from Svalbard and a form ofecological release (Leinaas 1999a). but a study designed to describe chironomid popu­ Fjellberg (1997) recently identified three col­ lation cycles revealed several new species (Hod­ lembolan species new to Svalbard which appear kinson et al. 1996a) and the current total is likely to be restricted to Nordaustlandet, Bonetogastrura to be a substantial underestimate. niva/is, Vertagopus arcticus and Anurida mar­ Hodkinson et al. (1996a) demonstrated the tem­ itima. perature dependency of chironomid emergence The Collembola fauna has been shown to be a on Svalbard and that in unusually warm years useful system by which to model general ecologi­ there was an initial wave of emergence early in cal theory (in particular metapopulation dynam­ summer with fewer individuals appearing in the ics and the influence of landscape heterogeneity second half. Chironomids form an important con­ and patchyness of population processes) (Hertz­ stituent ofthe diet ofsome arctic birds, for exam­ berg et al. 1994, 1998, Hertzberg 1997, Coulson ple the purple sandpiper (c. maritima) (Bengtson et al., in press). & Fjellberg 1975) and Hodkinson et al. (1996a) suggest that changes in the seasonal availability Class Insecta ofchironomids may have important consequences for breeding bird populations. The reported The class Insecta consists of 29 orders (Speight swarming behaviour ofone species ofchironomid, et al. 1999) but only nine orders are represented Smittia extrema, is unusual. The mating swarm on Svalbard (excluding the Ephemeroptera which is orientated about pale areas of ground set consists of one species of dubious validity; Table amongst the more uniform dark background. The 1) (Coulson & Refseth, in press). In common with males take flight on hearing the hum of the fe­ other arctic regions, Svalbard has a low number males wings as she takes off. Once copulation has of beetles and overwhelming preponderance of been occurred the female lands and the males fol­ Diptera species when compared with temperate low suit (Syrjiirniiki 1968). This is in contrast to latitudes. For example, 55% of the insect species the swarming flights of extended duration found on Svalbard are Diptera while the Coleoptera form more generally among the Chironomidae and only 8% ofthe insect fauna. Similarly, in the North seems to be an adaptation to the more open land­ American Arctic, the Diptera comprise 53% and scape and the problems of flight in the typically the Coleoptera 12% of the total species assem­ gusty wind conditions found in the open tundra. blage (Kevan & Danks 1986b). However, in the Syrjiimaki (1968) writes that he deduced the trig­ North American fauna (excluding the American ger to the flight of the males after observing their Arctic) the positions are reversed and the Diptera reaction to his humming an old Finnish folk tune. and Coleoptera form 19 and 33% of the total in­

50 Norw. 1. En/omol. 41, 41-63. 2000

Perhaps a rare example of the perfect fusion of Order Hymenoptera art and science. Five families ofHymenoptera are present on Sval­ There seem to be no records ofthe housefly Musca bard (Table I) with the Ichneumonidae forming domestica. Nevertheless, this species has a glo­ the most numerous family (16 species). One indi­ bal synanthropic distribution (Skidmore 1985) and vidual of an unknown species of cynipid that is it seems quite likely that the fly is present in ei­ probably an aphid parasitoid has been collected ther Longyearbyen or Barentsburg. The dipteran (Hodkinson et al. 1998). The sawfly Pontopristia parasites ofthe reindeer, the nose bot fly (Cephe­ amentorum is common on the tundra heath veg­ nemyia trompe), and the warble fly (Hypoderma etation where the larvae feed and pupate on the tarandi) are not recorded from Svalbard despite a leaves of Sa/ix polaris. Adults fly very close to palaearctic distribution which includes arctic the ground and seem to have two periods of diel Canada and Greenland (Danks 1981). Both these activity, between 10:00 and 12:00 and 14:00 to species have obligate one year life cycles, drop­ 15:00, on warm July days (pers. obs.). ping from the host as final instar larvae in May The ants (Formicidae) are notable by their ab­ and June. Subsequent summer temperatures must sence, but are generally rare in Arctic regions be high enough to permit pupation, adult emer­ (Kevan & Danks 1986b). Interestingly, neither gence and egg deposition (Nilssen 1997). Moreo­ species ofbumble bee (Bombus hyperboreous and ver, after emergence the imagines require air tem­ B. polaris: Apidae) present in Greenland (Danks peratures above lOoC in order to fly (Anderson & 1981) have been seen on Svalbard. Nilssen 1996). Infestation ofthe reindeer in North­ ern Norway is, at least in part, dependent on sum­ Order Co/eoptera mer temperatures (Nilssen 1997) and the short cool Svalbard summers may prevent colonisation Despite forming the largest insect order on a glo­ by these two parasites. There is one mosquito bal basis (Speight et al. 1999), the Coleoptera are present on Svalbard, Aedes (Ochlerotatus) nigri­ generally poorly represented inArctic regions (Ke­ pes. This species can be common in Longyear­ van & Danks 1986b). The pattern is repeated on byen and Brucebyen (Summerhayes & Elton Svalbard from where only 19 species of Coleo­ 1923, pers. obs.). ptera have been recorded from a total of231 spe­ cies ofinsect. Ofthese, one species (Oryzaephilus surinamensis) was collected from stored food­ Order Lepidoptera stuffs and is certainly an human introduction (Kan­ The Lepidoptera fauna consists of II species from gas 1967). The remaining 18 species are distrib­ seven families (Table I). However, it seems likely uted amongst nine families of which half belong that the majority are wind blown migrants and do to the Staphylinidae. not breed on the islands while one (Parnassius apo//o) is a dubious record. Only Apamea mai/­ Beetles are not common on Svalbard. Some of lardi and possibly Plute//a polaris are suspected the beetles appear highly restricted in distribution, of being resident (Kaisila 1973a, Alendal et al. three species ofstaphylinid are known from Sval­ 1980). Although the larvae of A. mai//ardi feed bard only from the Woodfjord region (Olophrum on the widely distributed tussock forming Gra­ boreale, Eucnecosum brachypterum and Oma/ium minae, the moth is only present in low densities casum) (Fjellberg 1983). It is possible that 0. and/or with a restricted distribution in western casum requires the higher soil temperatures that regions of Spitsbergen areas with an optimally occur in the vicinity ofthe warm springs (Fjellberg mild climate (Alendal et al. 1980, pers. obs.). All 1983). The staphylinids Atheta graminicola and species have a wide European distribution. Boreophi/a subplana are found in greatest num­ bers amongst the moist and luxuriant vegetation beneath the bird cliffs, for example the southerly facing cliffs at Kjrerstranda (Bf0ggerhalv0ya, Engelskbukta) and Grumant (lsfjord). Interest­

51 Coulson: Invertebrate fauna of the High Arctic archipelago of Svalbard

Kongsfjord A v Figure 3. A. Map of the Br0g­ gerhalv0ya. The collembolan M. arcllca is present in large Brs;;gger­ numbers under stones at the Engelskbukta base of both the Kjcerstranda halvs;;ya and Stuphallet bird cliffs. The predatory staphylinid beetle A. gramlnico/a is only found at B Kjcerstranda...... =escarp­ Sept. Oct. Nov. Dec. Jan. Feb. Mar. April May June ment, 6 o I', I , I I I I I B. Mean monthly temperatures 0_ recorded at a depth of 3cm ~ -5 1993/94. Kjcerstranda = dotted :::J line, Stuphallet solid line, '§ -10 = Q) (temperature data redrawn ~ -15 from Table 2, Coulson et al. Q) 1995). I- -20 ingly, even though the collembolan prey species are endemics, Acyrthosiphon svalbardicum and (M. arctica) is abundant, the beetles do not occur A. calvulus. Only A. svalbardicum has been de­ at the Stuphallet cliffs along the northern coast of scribed in detail (Strathdee et al. 1993a, Strathdee the Broggerhaloya (Figure 3). This is a region that & Bale 1995). The other species A. calvulus, has much lower winter and presumably summer Cinara abieticola and Pemphigus groenlandicus temperatures (Coulson et al. 1995, pers. obs.). The are only known from isolated records (Ossiannil­ distributions of these beetles may therefore be in sson 1958). While the host plant of A. calvulus part temperature limited. Hagvar (1971) points out (Poa arctica), is present on Svalbard, the hosts of that these species, and also RhynchaenusjIagellum the two other species are absent and the aphids and Simplocaria metallica, have been found only are likely to be random migrants (Ossiannilson where the vegetation is well developed. Micra­ 1958). Acyrthosiphon svalbardicum (host plant lymma marinum is probably restricted to beach Dryas octopetala) has a unique and genetically habitats (Hagvar 1971). determined abbreviated life cycle that is highly adapted to the short Arctic summer (Strathdee et Physiological studies on Amara quenseli indicate al. 1993a). Environmental cues such as short pho­ that the beetle on Svalbard has an elevated meta­ toperiod, crowding or poor host quality have no bolic rate compared to individuals from more effect on the life cycle and fail to trigger the pro­ southerly regions (Stromme et al. 1986). This is duction of alate or sexual morphs in contrast to interpreted as an adaptation to maintain activity the majority ofaphid species. Despite feeding on at the low environmental temperatures on a widespread host plant, the aphid only occurs in Svalbard. areas where the duration and mean temperature of the summer are sufficient to allow the insect to Order Hemiptera complete its annual life cycle. With the 'extra' Of the four species of Hemiptera that have been generation life history strategy, the insect seems recorded from Svalbard all are Aphididae and two ideally situated to take advantage ofglobal warm­

52

! L Norw. J Entomol. 47, 41-63. 2000 ing and increase both its density and range Orders Ephemeroptera, Trichoptera and (Strathdee et al. 1993a, 1995). This aphid was first Ne uroptera described from Vestpynten, Isfjord, by Ossiannils­ The insect orders Ephemeroptera and Trichoptera son (1958) but has since been collected from the are each represented by only one species, Lepto­ southern shore of Kongsfjord in the vicinity of phlebia vespertina and Apatania zonella, respec­ Ny-Alesund (Strathdee et al. 1993b) and more tively. Since 1. vespertina has only been recorded recently from Brucebyen (Billefjorden) in August once (J0rgensen & Eie 1993), is rare above the 1999 (pers. obs.). It is interesting to note that the tree line and would not be expected to be found aphid appears relatively widespread on the D. on Svalbard it is considered a dubious record (J.E. octopetala surrounding Brucebyen, but that Brittain, pers. comm.). It is unlikely that there are Summerhayes & Elton (1923) did not record the any species of Ephemeroptera present on Sval­ presence of the aphid despite extensive investi­ bard. In contrast, A. zonella is a typically arctic gation of the flora and invertebrate fauna of the species and the only caddisfly extending into the area. Whether the aphid has only recently colo­ High Arctic (Kevan & Danks 1986b). This spe­ nised Kapp Napier and Brucebyen is open to cies is probably uncommon on Svalbard being re­ speculation. stricted to regions where the climate is less harsh, for example Woodfjord (Decamps & Voisin 1971). Orders Anoplura and Mallophaga Early records often synonymise A. zonella (Tri­ The sucking lice, Anoplura, has two species on choptera) with Goniataulius arcticus (Neuropte­ Svalbard (Table 1), one, Echinophthirius horridus, ra). There are no valid species ofNeuroptera cur­ a parasite ofthe ringed seal (Phoca hispida), the rently recorded from Svalbard. other, Antarctophthirus tricechi, from the walrus (Odobenus rosmarus) (Kaisila 1973b). A recent Phylum Crustacea revision of the literature concerning the Mallo­ No terrestrial Crustacea occur on Svalbard (Coul­ phaga of Svalbard concluded that there were 35 son & Refseth, in press). However, there are four species from two families parasitising the birds classes offreshwater Crustacea present; Branchio­ (Mehi et al. 1982). poda, Copepoda, Ostracoda and Malacostraca (Table 1). The cladoceran Chydorus sphaericus Order Siphonaptera is perhaps the most common crustacean and is The seabirds and geese are hosts to just two spe­ found in both the pelagic and litteral zones cies of Siphonaptera. Miotenopsylla arctica arc­ (J0rgensen & Eie 1993). The crustacean that has tica is found in the nests of the kittywake (Rissa attracted the greatest attention however, is the tridactyla) while the second flea, Ceratophyllus water flea, Daphnia pulex (Cladocera). This spe­ vagabundus vagabundus parasitises the pink cies has a circumarctic distribution and detailed footed geese (Anser brachyrhynchus), barnacle genetic studies have shown D. pulex to be a spe­ geese (E. leucopsis) and gulls (Mehl1992). Fleas cies complex with two major lineage groups and were present on 5% of the adult birds but nest­ two zones of overlap, one in N. Europe and the lings had a higher incidence of infection, up to other in Canada (Hobrek & Weider 1999). As with 60% in young kittywakes and 12% in young puf­ the tardigrades (Pugh & McInnes 1998), the fins (Fratercula arctica). No fleas were found in spread of the various Daphnia clones appears to the nests of fulmars (Fulmaris glacialis), storm generally eastwards but for unknown reasons petrels (Hydrobates pelagicus) or arctic skuas (Roen 1981, Hobrek & Weider 1999). The authors (Stecorarius parasiticus) although the number of contend that, with as much clonal diversity nests searched was low. In northern regions on amongst high arctic Daphnia as in the temperate Svalbard it is only M arctica arctica that is present zone, the view ofthe arctic as a homogeneous bio­ despite both species having a circumarctic distri­ me must be re-evaluated. Two phenotypic morphs bution (Mehl 1992). ofD. pulex occur on Svalbard, a melanic dark form and a paler non-melanised form (Hessen 1996).

53 Coulson: Invertebrate fauna of the High Arctic archipelago of Svalbard

While the melanised form appears to have a slower Meijering 1985). The copepod Limnocalanus ma­ growth rate, it also has increased tolerance to high crurus (Copepoda: Calanoida) is present over a UV-B light levels. Itis possible that the frequency great part ofEurope but may be a relict arctic spe­ ofmelanic/non-melanic clones may be explained cies since it has a higher rate ofdevelopment than as a trade-offbetween the metabolic costs ofmela­ other copepods and temperatures over l6°C are nin production and UV-B protection (Hessen lethal to the eggs (Jmgensen and Eie 1993). One 1996). Despite having altered their developmen­ copopod parasitising the arctic charr is also pre­ tal rate so as to be able to complete their annual sent, Salmonicola edwardsii (Siphonostomatoida) life cycle within the short arctic summer, the distri­ (Kennedy 1978, Sobecka & Piasecki 1993). bution ofD. pulex on Bear Island remains restric­ ted to small permanent waters since temporary DISCUSSION ponds are too ephemeral for the crustacean to com­ plete its life cycle and fish predators prevent colo­ Fauna nisation of the larger lakes (Meijering & Jacobi The species paucity of the invertebrate fauna of 1981). Macrothrix hirsuticornis (Cladocera) ap­ Svalbard is striking when considered against other pears to be partially pre-adapted to the arctic con­ arctic regions such as arctic Canada and Green­ ditions since the temperature responses of indi­ land. It is difficult to directly compare species di­ viduals from Svalbard are identical to those from versity between regions due to differences in the Europe (Meijering 1979). Jfilrgensen & Eie (1993) current knowledge ofspecies assemblages and the consider that the species is only present in shal­ status of taxanomical revision. However, by us­ low lakes that freeze to bottom in winter and that ing well known groups, for example the Araneae, have an average July/August water temperature some comparisons can be made. There are 18 spe­ of6°C. Without such temperatures the animal has cies of spider from three families known from insufficient physiological time to produce the Svalbard but the total on Greenland stands at 43 overwintering eggs. Lepidurus arcticus (Branchio­ species from ten families (Danks 1981) (Table 3). poda: Notostraca) has a fairly widespread distri­ The Greenland invertebrate fauna contains II or­ bution throughout the archipelago and is indica­ ders ofInsecta while just nine are found on Sval­ tive ofcold clean freshwaters. This crustacean can bard (Danks 1981) (Table 3). Furthermore, 23 spe­ form a major food source for the arctic charr cies of Cladocera were recorded from just one (Salvelinus arcticus) and presence or absence can region of Greenland (Roen 1997) compared with affect charr morph dynamics (Klemetsen et al. only eight from the whole of Svalbard (Table 3). 1985). Moreover, the animal disturbs the lake beds It is likely that this lack of species diversity on while searching for the food, mixing the detritus Svalbard is due to three factors operating indepen­ with the water column and so releasing nutrients dently and in combination: i) the remoteness of for use by the plankton fauna (Jmgensen & Eie Svalbard from the nearest mainland, ii) the cli­ 1993). mate and iii) the young age ofthe ecosystem. Since the closest mainland is ca. 1000 km distant, im­ The four species ofCyclopidae, for example Cy­ migration for many non-aerial species such as clops abyssorum, are widespread throughout Sval­ Collembola is probably infrequent. Despite the bard including Bear Island. As with the Cladocera, high arctic latitude of the archipelago, the winter the Cyclopidae fauna is also comprised of eurye­ climate ofSvalbard is mild with average monthly cious species (Koch & Meijering 1985). The Cy­ winter air temperatures being only a little below ­ clopidae potentially have the ability to overwinter IOOC (Ffilrlandet al. 1997b). Moreover, the sum­ as either adults or any copepodid stage. However, mer is also short and cool, even on the warmer the stage employed depends on the habitat type. west coast the soil is typically frozen for over 300 In cold ponds where development is delayed it is days each year (Coulson et al. 1995). An absence the young stages that overwinter, while in warmer of intermediate island groups to act as 'stepping ponds the animals have sufficient physiological stones' between the mainland species source and time to develop into the 4th or 51h stages (Koch & Svalbard will both reduce the rate of species im­

54 Norw. J Entomo/. 41, 41-63. 2000 migration and also removes the opportunity for is either descended from a fauna that survived the gradual species adjustment to the Svalbard envi­ glacial period in-situ in unglaciated refugia, for ronment via the colonisation of intervening is­ example, nunataks (unglaciated mountain tops lands. Any species that is to become established above the ice sheets), or the glaciation 'wiped the on Svalbard must possess a physiology and a life slate clean' and the fauna consists of those spe­ history which immediately enables survival ofthe cies that subsequently spread out from more south­ characteristic seasons ofSvalbard. Ecosystem age erly unglaciated latitudes following the retreat of may also have a role in limiting species diversity the ice sheets (the 'tabula rasa' hypothesis) (Ives on Svalbard. While the insect fauna of Hawaii 1974). stands at 7862 species, no less than 5237 ofwhich are endemic, there are only four endemics from Dispersal of the invertebrate fauna to Svalbard Greenland from an unknown number of insects There are four means by which invertebrates from (Sadler 1999) (but 255 species are cited in Danks the mainland can colonise an island, via; i) wind (1981) which is surely an underestimate of the currents (and frontal systems), ii) oceanic currents species total). Sadler (1999) considers that a prime (drift wood, ice rafting etc.), iii) animal vectors factor in limiting species diversity among the and iv) human introductions (Whittaker et a1. North Atlantic islands has been the 'largescale and 1989, Christiansen & Bellinger 1994, Bergersen periodic disturbance by Quaternary icesheets' . 1995).

Origin of the invertebrate species on i) wind currents. Strong low pressure systems Svalbard moving over Svalbard from the Atlantic result in powerful south westerly winds. Lokki et a1. (1978) During the WisconsinianIWiirm event (170,000­ records the arrival oflarge numbers of the butter­ 10,000 years RP.) Svalbard was heavily glaciated, flies P xylostella and one individual of rilnessa deglaciation occurring in during the Younger cardui on Svalbard after strong south-easterly Dryas (11,000 to 10,000 years RP.) (Salvigsen & winds during a severe storm (wind velocities of Osterholm 1982). The current fauna of Svalbard

Table 3. Number of freshwater and terrestrial invertebrate families on Svalbard and Green­ land (Greenland data from Danks (1981) and Holland (1985). a includes the Psylloidea, Hodkinson (1997), b plus one extra possible family, 'dubious record, J0rgensen & Eie (1993), dincluding the dubious ephemeropteran).

Class Order Total number of families Svalbard Greenland Arachnida Araneae 3 10 Collembola 10 9 Insecta Anoplura 1 3 Hemiptera 1 sa Mallophaga 2 3 Dictuoptera 0 1 Dermaptera 0 1 Coleoptera 10 5b Diptera 21 22 Lepidoptera 7 11 Hymenoptera 5 4 Siphonaptera 1 3 Trichoptera 1 1 Ephemeroptera l' 0 Total 63d 78

55 eoulson: Invertebrate fauna of the High Arctic archipelago of Svalbard

24 m sec-Ion Spitsbergen), a 1000 km journey this manner. However, Seyd (1979) points out that that took 24 h. That the airborne route may be the oribatid nymphs have been found in moss carpets most important means of arrival on Svalbard is from the middle ofthe crater. It is hard to imagine also indicated by the number of Diptera species how they arrived by hydrochorous drift and Seyd on Svalbard. With a total of 128 species this or­ (1979) considers wind transport is more likely in der comes third after the Rhizopoda and Rotifera this particular case. Large amounts of driftwood (where wind dispersal is also considered to be im­ originating in the river estuaries of north Russia portant) in terms of species richness. For exam­ arrives on the Spitsbergen coast (Johansen 1999). ple, Elton (1926) notes the occurrence ofSyrphus The recent finding that some Arctic soil inverte­ ribesii (Diptera: Syrphidae) on Svalbard after a brates can survive several years at sub-zero tempe­ large depression had moved over Svalbard. The ratures (Coulson & Birkemoe submitted) opens airborne route is also favoured by Holm (1958) the possibility ofspecies being transported to Sval­ as an explanation as to the means ofarrival ofthe bard on such wood while frozen in the sea ice. spider fauna. Holm (1958) listed just 15 species iii) animal transport. Vertebrate animals may have ofspider from Svalbard. At the same time 21 spe­ been responsible for the introduction ofmany in­ cies were known from the Scoresby sound region vertebrate species. Collembola may be transported of Greenland alone, an area that has a similar cli­ to remote islands in mud on the feet of birds, al­ mate. Holm (1958) concludes that aerial disper­ though direct evidence ofthis is lacking (Christi­ sal is the method by which the majority of the ansen & Bellinger 1994). A total of 163 species spiders have colonised Spitsbergen since this strat­ ofbird are recorded for Svalbard, 41 ofwhich are egy is well documented for many Araneae. In sup­ known to breed on the islands, the remainder be­ port of this he cites the fact that it is the larger ing vagrants (Mehlum 1990). Many bird species spider species present in Greenland, and presum­ travel large distances. For example, the barnacle ably those less likely to be swept up into the aerial goose (B. leucopsis), the entire Svalbard popula­ plankton, that are absent on Svalbard. Wind is also tion of which overwinters at Caerlaverock on the the most probable means of dispersal ofboth the Solway Firth in Scotland, a distance of 3000 km Tardigrada and the Rotifera, a method that may from Svalbard (Owen 1990). During these move­ include temporary refuge in cryoconite holes (de ments the birds make use of numerous staging Smet 1993, de Smet & van Rompu 1994, Pugh & posts on the Norwegian mainland and coastal is­ McInnes 1998). From an analysis of the circum­ lands with the possibility to pick up and/or ex­ arctic Tardigarde species assemblages, Pugh & change hitchhiking species. However, many bird Mclnnes (1998) concluded that few, if any, tardi­ species spend time at sea before coming ashore grades survived the Pleistocene glaciation in-situ, and the salt water would kill many terrestrial or but that the arctic has been recolonised during the freshwater invertebrates hitchhiking on the ani­ Holocene by a common non-arctic fauna origina­ mals body. Other birds arrive before the land has ting in N. America and spread by the prevailing cleared ofsnow and the majority ofeggs/animals westerly winds during the recent Holocene. deposited under such conditions will be washed ii) oceanic currents. While it seems possible that away during the brief period of snow melt (de species have arrived on Svalbard due to ocean cur­ Smet 1993). rents, most probably hitching a ride on driftwood, Many of the parasites of the vertebrates are po­ it is difficult to find much direct evidence for it. tentially highly mobile by virtue of their hosts. Nevertheless, the beach-dwelling beetle, M. mari­ The global polar bear population tends to be di­ num, probably colonised Svalbard in this manner vided into sub-populations but there is a degree (Hagvar 1971). Lindroth et al. (1973) demon­ of intermixing. Tagged bears from the Svalbard strated that many oribatid mites and Collembola population have, for example, been recaptured in could survive over a week on floating grass tus­ Greenland (Wiig 1995) while Svalbard bears have socks and suggest that many of the Collembola also been identified from the Russian Arctic (Fogg and mites that have colonised Surtsey arrived by

56 Norw. J. Enlomo/. 47, 41-63. 2000

1998). Such migrations obviously have implica­ 1998). Pugh & Mclnnes (1998) found no evidence tions for the spread of organisms parasitising or for glacial refugia amongst extant tardigrade spe­ travelling on the bears. The voles have recently cies assemblages, a group with extreme desicca­ been shown to be infected with the cestode E. mul­ tion and cold tolerance (Somme 1996). Opalinski ti/ocularis. Before the vole was accidentally intro­ & Klekowski (1992) studied the metabolic tem­ duced to Svalbard early this century there was no perature relations of several freshwater inverte­ suitable intermediate host for the parasite on the brate taxa from Svalbard. As few species displayed islands. It is likely therefore that the parasite trav­ physiological temperature independence or indi­ elled to Svalbard with the voles which originated cations ofmetabolic cold compensation they con­ from the Leningrad region of Russia (Fredga et cluded that the ecosystem was ecologically young al. 1990). In addition the voles are host to two and that there was a predominance of pioneering species ofparasitic mites, Laelaps hi/aris and Hae­ ubiquitous species. However, an unusual form of mogamasus ambulans, that were probably intro­ refugia on Greenland has been proposed by Ber­ duced to Svalbard along with the rodents. gersen (1995). While considering the endemic in­ vertebrate fauna of Greenland, Bergersen (1995) iv) human introductions. There are few reliable noted that the majority ofthese species were aqua­ reports of invertebrates that have been acciden­ tic or moisture-loving and occur in South and tally introduced due to human activity, with the South Western Greenland. Bergersen (1995) sug­ exception of the parasites associated with the in­ gests that the warm springs present in this region troduced mammals, and none seem to have be­ gave additional buffering to that often provided come established. Summerhayes & Elton (1928) by water (Danks 1992) to climatic extremes dur­ report seeing the fly Calliphora vicina (Diptera: ing the glaciations that denuded the fauna of Calliphoridae) on board a ship in 1924. This is Northern Greenland, there being no endemic in­ the only record ofthis species and it had probably vertebrate or plant from this region. It is addition­ arrived on board the ship. Moreover, all observa­ ally suggested that the mildly radioactive waters tions ofPhormia terrae-novae (Diptera: Callipho­ ofthese springs may have accelerated the rate of ridae) have been in the vicinity of human dwell­ speciation. A detailed investigation of the fauna ings (Nourteva 1967) and it seems likely that this associated with the Bockfjorden warm springs on species is synanthropic on Svalbard. Kangas Svalbard might prove interesting. Survival in ice­ (1967) found one dead adult ofthe beetle, Oryzae­ free regions was also considered the best expla­ phi/us surinamensis (Coleoptera: Silvanidae) in nation of the current discontinuous distribution a packet of biscuits in a field hut. This is a com­ of the mite Calyptozetes sarekensis by Seyd mon pest of stored foodstuffs and is unlikely to (1979). Seyd (1979) suggested that C. sarekensis be resident. is an ancient species dating from the time of the Laurasian continent (60 million years BP) and that Glacial refugia: the nunatak hypothesis a once homogeneous range was cleaved by subse­ The nunatak glacial refuge hypothesis for arctic quent ice sheets. species has received considerable attention (Ives 1974, Dahl 1987). While nunataks certainly do Conclusion form refugia for Arctic and Antarctic fauna and In conclusion, the invertebrate fauna of Svalbard flora (Roen 1981, Bostrom 1997, Cocks et al. is characterised by its paucity compared to other 1998, Strong 1999), their importance in the sur­ Arctic regions. This is likely to be due to a com­ vival offaunal and floral assemblages in high arc­ bination of the remoteness of the islands and the tic regions during the Pleistocene glaciation is ecologically young age of the ecosystem. The unclear and disputed (Dalh 1987, Andersen 1988, number ofspecies described from the archipelago Birks 1996, Bocher 1997, Sadler 1998). One prob­ currently stands at 1040 but this number will lem is that even if ice-free refugia existed they change as new species are documented and clas­ would have had an extreme arctic climate (Sadler sifications revised. The majority of the fauna con­

57 Coulson: Invertebrate fauna of the High Arctic archipelago of Svalbard sists of circumpolar species and there are rela­ Beyens, L., Chardez, D., De Landtsheer, R., with col­ tively few endemics. Although much ofthe fauna laboration of De-Bock, P. & Jacques, E. 1986a. Tes­ has closer affinities with that of North America tate amoebae populations from moss and lichen than Europe (Loof 1971, Pugh & McInnes 1998, habitats in the Arctic. Polar BioI. 5, 165-173. Beyens, L., Chardez, D., De Landtsheer, R., with col­ Hobrek & Weider 1999) some groups, for exam­ laboration of Baere, 0. 1986b. Testate amoebae ple the Lepidoptera, are comprised mainly ofEu­ communities from aquatic habitats in the Arctic. ropean species. The focus during the last 10 years Polar BioI. 6, 197-205. has moved away from reports dealing with the Beyens, L. & Chardez, D. 1995.An annotated list of presence of invertebrate species towards articles testate amoebae observed in the Arctic between the using manipulative field and laboratory experi­ longitudes 27°E and 168°W Arch. Protistenkd. 146, ments to explore the ecology and physiology of 219-233. the fauna. Beyens, L. & Chardez, D. 1997. New testate amoebae taxa from the polar regions. Acta Protozool. 36, 137­ 142. Acknowledgements. The constructive comments and Birkemoe, T. 1995. Population dynamics ofEnchytrae­ A suggestions ofL. S0mme, Fjellberg andA.C. Nilssen idae at the Arctic tundra at Spitsbergen, Svalbard. were very much appreciated. Newslett. Enchytraeidae 4, 42-52. Birkemoe, T. & S0mme, L.S. 1998. Population dynam­ References ics of two collembolan species in an arctic tundra. Alendal, E. & Helle, O. 1983. Helminth parasites of Pedobiologia 42, 131-145. muskoxen Ovibos moschatus in Norway including Birkemoe, T. & Leinaas, H.P. 1999. Reproductive bi­ Spitsbergen and Sweden with a synopsis of para­ ology of the arctic collembolan Hypogastrura sites reported from this host. Fauna norv. Ser. A 4, tullbergi. Ecography 22,31-39. 41-52. Birkemoe, T., Coulson, S.l & S0mme, L. (submitted). Alendal, E., Andersen, T. & Fjeldsa, A. 1980. New Species characteristics, life cycles and population records of Apamea maillardi (Geyer, 1932) (Lepi­ dynamics of enchytraeids (Oligochaeta) from the doptera: Noctuidae) in Adventdalen, Spitsbergen. high Arctic. Fauna norv. Ser. B 27, 78. Birks, H.J.B. 1996. Statistical approaches to interpret­ Amren, H. 1964. Ecological and taxonomical studies ing diversity patterns in the Norwegian mountain on zooplankton from Spitsbergen. Zool. Bidr. Upps. flora. Ecography 19,332-340. 36, 209-277. Block, W, Webb, N.R., Coulson, S., Hodkinson, LD. Andersen, J. 1988. Post glacial immigration of plants & Worland, M.R. 1994. Thermal adaptation in the and animals in Scandinavia. Fauna (Oslo) 41, I­ arctic collembolan Onychiurus arcticus (Tullberg). ll. J. Insect Physiol. 40,715-722. Anderson, IR. & Nilssen, A.C. 1996. Trapping oestrid Bocher,l 1997. A history ofthe insect fauna ofGreen­ parasites ofreindeer: The response of Cephenemyia land. In Ashworth, A, Buckland, P.c. and Sadler, trompe and Hypoderma tarandi to baited traps. IP. (eds.) Studies in Quaternary entomology: an Med. Vet. Ent. 10,337-346. inordinate fondness ofinsects. Quaternary Proceed­ Balik, V. 1994. On the soil testate amoebae fauna (Pro­ ings 5. Wiley, Chichester. tozoa: Rhizopoda) of the Spitsbergen Islands Bostriim, S. 1997. Chiloplectus masleni sp. novo & vari­ (Svalbard). Arch. Protistenkd. 144,365-372. ability in populations ofPlectus acuminatus Bastian Bengtson, S.A., Fjellberg, A. and Solh0Y, T. 1974. 1865 (Nematoda: Plectidae) from the nunatak Abundance of tundra arthropods in Spitsbergen, Basen, Vestfjella, Dronning Maud Land, East Ant­ Norway. Ent. Scand. 5, 137-142. arctica. Polar BioI. 17,74-80. Bengtson, S.A. & Fjellberg, A. 1975. Summer food of Byzova, IB., Uvarov, AV. & Petrova, A.D. 1995. Sea­ the purple sandpiper Calidris maritima in sonal changes in communities of soil invertebrates Spitsbergen, Norway. Astarte, 8, 1-6. in tundra ecosystems of Hornsund, Spitsbergen. Bengtson, SA, Fjellberg,A. & Solh0Y, T. 1975. Amara Polar Res. 16,245-266. quenselii Schu. (Coleoptera: Carabidae) new to Chardez, D. & Beyens, L., with the collaboration of Svalbard, Norway. Norw. 1 Entomol. 22, 81-82. De Bock, P. 1987. Arcella ovaliformis sp. novo a Bergersen, R. 1995. Is Greenland a zoogeographical new testate amoeba from Edge0ya, a high Arctic unit of its own? 1 Biogeogr. 22, 1-6. island. Arch. Protistenkd. 134,297-301. Christiansen, K. & Bellinger, P. 1994. Biogeography

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61 Coulson: Invertebrate fauna of the High Arctic archipelago of Svalbard

Roen, D. 1997. The Cladocera ofthe Godthabfjord area, Strathdee, A.T., Bale, 1.S., Block, WC., Webb, N.R., S.W Greenland. Hydrobiologia 360, 75-78. Hodkinson, LD. & Coulson, S.J. 1993a. Extreme Rompu van, E.A & Smet de, WHo 1996. Freshwater adaptive life cycle in a High Arctic aphid, Acyrtho­ tardigrades from Hopen, Svalbard (76°31 'N). Fauna siphon svalbardicum. Ecol. Entomol. 18,254-258. norv. Ser. A 17, 1-9. Strathdee, AT., Bale, 1.S., Hodkinson, I.D., Block, W, Sadler,1.P. 1998. 'Is Greenland a zoogeographic unit?' Webb, N.R. & Coulson, S.1. 1993b. Identification A response to Bergersen. 1. Biogeogr. 25, 399-403. of three previously unknown morphs ofAcyrthosi­ Sadler, 1.P. 1999. Biodiversity on oceanic islands: a phon svalbardicum Heikinheimo (Hemiptera: palaeoecological assessment. 1. Biogeogr. 26, 75­ Aphididae) on Spitsbergen. Ent. Scand. 24, 43-47. 87. Strathdee, AT., Bale, 1.S., Strathdee, P.C., Block, WC., Sagerup, K., Gabrielsen, G.W, Skorping, A. & Skaare Coulson, S.1. & Webb, N.R, 1995. Climatic sever­ 1.D. 1998. Association between polychlorinated ity and the response to temperature elevation of biphenyl (PCB) concentrations nematodes in glau­ arctic aphids. Global Change BioI. I, 23-28. cous gulls, Lams hyperboreous, from Bear Island. Strathdee, A.T. & Bale, 1.S. 1995. Factors limiting the Organohalogen Compounds 39, 449-451. distribution ofAcyrthosiphon svalbardicum (Hemi­ Salvigsen, O. & 0sterholm, H. 1982. Radiocarbon dated ptera: Aphididae) on Spitsbergen. Polar BioI. 15, raised beaches and glacial history of the northern 375-380. coast of Spitsbergen, Svalbard. Polar Res. 1, 97­ Strong, WL. 1999. Mountain Park area: a plant refu­ 115. gium in the Canadian Rocky mountains? J. Bio­ Seyd, E.L. 1979. The evolution and distribution of geog. 26, 413-423. Calyptozetes sarekensis (Acari: Oribatei). BioI. 1. Strl'Jmme, J.A. 1989. Metabolic rates ofalpine and High Linn. Soc. 12, 1-18. Arctic Amara quenseli (Col., Carabidae) beetles. Skidmore, P. 1985. The biology ofthe Muscidae of the Fauna norv. Ser. B 36, 57-58. world. 550 pp. W Junk, Dordrecht. Summerhayes, VS. & Elton, C.S. 1923. Contributions Smet de, WHo 1993. Report on rotifers from Barents­ to the ecology of Spitsbergen and Bear Island. 1. l'Jya, Svalbard (78°30'N). Fauna norv. Ser. A 14, 1­ Ecol. 11,214-286. 26. Summerhayes, VS. & Elton, C.S. 1928. Further con­ Smet de, WHo & Rompu van, E.A 1994 -Rotifera and tributions to the ecology of Spitsbergen. 1. Ecol. Tardigrada from some Cryoconite holes on a Spits­ 16, 193-268. bergen (Svalbard) glacier. Belg. 1. Zool. 124, 27­ Sustr, T. & Block, W, 1998. Temperature dependence 37. and acclimatory response of amylase in the high Sobecka, E. & Piasecki, W 1993. Parasitic fauna of arctic springtail Onychiurus arcticus (Tullberg) Arctic charr, Salvelinus alpinus (L., 1758) from the compared with the temperate species Protaphomra Hornsund region (Spitsbergen). Acta lchylogica et armata (Tullberg). 1. Insect Physiol. 10, 991-999. Piscatoria, 23(supplement), 99-106. S)'Ijiimliki, S. 1968. A peculiar swarming mechanism Solem, J.O. & Sendstad, E. 1978. Diversity in diel pe­ of an arctic chironomid (Diptera) at Spitzbergen. riodicity ofCollembola communities at Spitsbergen Ann. zool. fenn. 5,151-152. Svalbard Norway. Norw. 1. Entomol. 25, 9-14. Tambs-Lyche, H. 1967. Notes on the distribution of Sl'Jmme, L. 1979, Insect life on Svalbard. Fauna (Oslo) some Arctic spiders. Astarte 28, 1-13. 32,137-144. Thor, S. 1930. Beitrage zur Kenntnis der invertebraten Sl'Jmme, L. 1993, The terrestrial arthropod fauna of Fauna von Svalbard. 155 pp. Skrifter om Svalbard Svalbard. Arctic Insect News 4,2-4. og Ishavet 27. Sl'Jmme, L. 1996. Anyhydrobiosis and cold tolerance Webb, N.R., Coulson, SJ., Hodkinson, I.D., Block, in tardigrades. Eur. 1. Entomol. 93, 349-357. W, Bale, 1.S. & Strathdee, A.T. 1998. The effects Sl'Jmme, L. & Birkemoe, T. 1997. Cold tolerance and of experimental temperature elevation on popula­ dehydration in Enchytraeidae from Svalbard. J. tions ofcryptiostigmatic mites in High Arctic soils. Comp. Physiol. B 167,264-269. Pedobiologia 42, 298-308. Sl'Jmme, L. & Birkemoe, T. 1999. Demography and Whittaker, R.1., Bush, M.B. & Richards, K. 1989. Plant population densities of Folsomia quadrioculata recolonisation and vegetation succession on the (Collembola, Isotomidae) on Spitsbergen. Norw. 1. Krakatau island, Indonesia. Ecol. Monographs 59, Entomol. 46, 35-45. 59-123. Speight, M.R., Hunter, M.D. & Watt, AD. 1999. Ecol­ Wiig, O. 1995. Distribution of polar bears (Ursus ogy of insects. Concepts and applications. 350 pp. maritimus) in the Svalbard area. J. ZooI. 237, 515­ Blackwell Science Ltd., Oxford.. 529.

62 Narw. 1. Enlamal. 41, 41-63. 2000

Worland, M.R. 1996. The relationship between water Haplophilus subterraneus (Shaw, content and cold tolerance in the arctic collembolan 1789), a centipede (Chilopoda, Onychiurus arcticus (Collembola: Onychiuridae). Eur. 1. Entomol. 93,341-348. Geophilomorpha) new to Norway Worland M.R., Grubor-Lajsic, G. & Montiel, P.O. 1998. Partial desiccation induced by sub-zero tempera­ Kjell Magne Olsen tures as a component of the suyrvival strategy of the Arctic collembolan Onychiurus arcticus (Tull­ berg). 1. Insect Physiol. 44, 211-219. Olsen, K.M. 2000. Haplophilus subterraneus (Shaw, Wiithrich, C. 1989. Die Bodenfauna in der Arctischen 1789), a centipede (Chilopoda, Geophilomorpha) new Umwelt des Kongsfjords (Spitzbergen) - Versuch to Norway. Norw. 1. Entomol. 47, 63-64. einer intergrativen Betrachtung eines Okosystems. Pp. 133. Basler Beitrage zur Physiogeographie, Four specimens of Haplophilus subterraneus (Shaw, Basel, Heft 12, 1789) were collected in the close vicinity of the com­ Wiithrich, C. 1991. Stofftransporte Meer-Land: Vogel­ post heap of the Botanical Garden, University of Oslo klifftundren und ornithogene Boden. Stuttgarter at TlIJyen, Oslo in 1992 and 1995. Adult specimens of Geographische Studien Bd. 117, 177-192. both sexes were present in both years, indicating a vi­ Wiithrich, C. 1992. Landschaftsokologische Umwelt­ able population at the locality. forschung: Beitrage zu den Wechselwirkungen zwischen biotischen und abiotischen Faktoren im Key words: Haplophilus subterraneus, Chilopoda hocharktischen Okosystem (Spitzbergen). Die Erde 122,335-352. Kjell Magne Olsen. Skarvelokka, N-4818 Fa:rvik, Nor­ way. Received 18 January 2000, accepted 15 March 2000 INTRODUCTION Haplophilus subterraneus (Shaw, 1789) is an ex­ clusively synanthropic centipede in the Nordic countries. The first find in the area was made in Rosenbergs Hage, a botanical garden in Copen­ hagen, Denmark (Bergs0e & Meinert 1866). Large numbers were present. Later, several other Dan­ ish localities has been added to the list (Enghoff 1973, 1983), one of these is an indoor find. In Sweden, the first specimens were found in a botanical garden in Visby, Gotland in 1913 (Porat 1913). Four males and 11 females were collected from underneath wooden flowerpots. Four specimens (1 juv., 1 0,299) were collected from greenhouses in southern Finland in 1947 and 1948 (Palmen 1948). H. subterraneus can be separated from the other Norwegian species of the order Geophilomorpha by this combination ofcharacteristics: coxal pores distributed over the whole surface ofthe hindmost coxa (Figure 1); head broader than long; more than 70 pairs oflegs (Eason 1964).

63 Short communication

H. SUBTERRANEUS IN NORWAY mainly carnivorous, and the area is rich in suit­ able invertebrate prey. H. subterraneus is also The compost heap at the Botanical Garden and known to occasionally feed on plant tissues Museum, University ofOslo at T0yen, Oslo (EIS (Thompson & Sankey 1961, Eason 1964). As this 28) was visited twice, on 30 May and 30 August is a more southern species, the relatively high win­ in 1992. The first visit revealed a male, approxi­ ter temperature inside the compost (due to fermen­ mately 36 mm long as preserved in alcohol, and tation) may provide the condition necessary for it with 79 pairs of legs. The second time one male, to survive the winter at the locality. 42 mm long and with 79 pairs of legs, and one female, 41 mm long and with 81 pairs of legs, Acknowledgements. I wish to thank Arvid Bach for were found. The same place was visited again on correcting my English. 4 May 1995. This time a female, 47 mm long and with 83 pairs oflegs was found. The preservation References of the animals result in a shortening of the body Bergsee, V. & Meinert, F. 1866. Danmarks geophiler. length, as 50-60 (70) mm are normal lengths of Naturhistorisk Tidsskrift. 3. Ra:kke IV (l), 81-108. fresh specimens (Eason 1964). Cloudsley-Thompson, IL. & Sankey, 1. 1961. Land invertebrates. A guide to British worms, molluscs The locality is highly synanthropic, and plants are and arthropods (Excluding insects). Methuen & Co. regularly introduced to the garden and the close­ Ltd., London. by greenhouses from abroad (R. Elven pers. Eason, E.H. 1964. Centipedes of the British Isles. 294 comm.). pp. Frederick Warne , London. Enghoff, H. 1973. Diplopoda and Chilopoda from sub­ Members of the order Geophilomorpha usually urban localities around Copenhagen. Vid. Medd. take about three years to reach maturity (Eason Dansk nat.hist. For. 136,43-48. 1964). The fact that adult specimens ofboth sexes Enghoff, H. 1983. Oversigt over skolopendrenes udbre­ were found suggests that a maintainable popula­ delse i Danmark (Chilopoda). Entomo1. Medd. 50, tion is inhabiting the area. Geophilomorphs are 1-6. Palmen, E. 1948. The Chilopoda ofeastern Fennoscan­ dia. Ann. Zoo1. Soc. Zoo1.-Bot. Fenn. "Vanamo" 13 (4), 1-46. Porat, C.O.v. 1913. En fcir Sverige ny myriopod. Ent. Tidskr. 34 (2-4), 176-178.

Received 15 September 1999, accepted 30 November 1999

Figure 1. Hap/ophi/us subterraneus (Shaw, 1789). Ventral view of posterior end of male (from Eason 1964) .

64 Norw. J. Entomol. 41, 65-11. 2000

New records of Norwegian Scatopsidae (Diptera)

Jean-Paul Haenni & Uta Greve

Haenni, 1.- P. & Greve, L. 2000. New records ofNorwegian Scatopsidae (Diptera). Norw. 1. Entomol. 47,65-71.

Distributional data are given for 26 species of Scatopsidae, 13 of which are new records for the fauna of Norway, i. e. Ectaetia christii Rotheray & Horsfield, E. clavipes (Loew), E. platyscelis (Loew), Thripomorpha bifida (Zilahi-Sebess), Apiloscatopse bifilata (Haliday), A.flavocincta (Duda), Apiloscatopse scutellata (Loew), Colobostema obscuritarse (Strobl), C. triste (Zetterstedt), Holoplagia bullata (Edwards), H. transversalis (Loew), Swammerdamella acuta Cook and S. adercotris Cook. A total of 31 species are recorded from Norway and the faunistics of the family is discussed.

Keywords: Scatopsidae, Diptera, Norway.

Jean-Paul Haenni, Museum d'histoire naturelle, rue des Terreaux 14, CH-2000 Neuchatel, Switzer­ land. Lita Greve, Zoological Museum, University ofBergen, Museplass 3, N-5007 Bergen, Norway.

INTRODUCTION parts of Norway in the course of various studies not devoted to this family. Identifications have Study of Norwegian Scatopsidae has been much been partly made by LG but most (including all neglected till recently. Scanty literature include difficult cases) have been performed or checked older records as Zetterstedt (1838, 1850), Siebke by JPH, who is responsible for them. All the ma­ (1877), Schoyen (1889) and Lundstrom (1913), terial is deposited in the collections of the Zoo­ as well as recent studies by Cook (1974) and logical Museum of the University of Bergen Andersson (1978, 1982). (ZMB), except for few specimens retained in the Our first faunistic article about Scatopsidae from collection ofJPH in Museum d'histoire naturelle Norway (Haenni & Greve 1995) summarised the de Neuchatel, Switzerland (MHNN). knowledge of the family in this country, record­ The following abbreviations are used for meth­ ing 19 species, ofwhich 9 were new records. Since ods of capture: B = beating; BT = Barber trap; then, new material has been gathered and stud­ CN = car-netting; F = fogging; LT = light-trap; ied, allowing the discovery ofseveral till now un­ MT = Malaise trap; N = net; WT = window trap recorded species and extending the known range (collision trap); YT = yellow tray. of some formerly recorded species. The present article follows the format of our preceding paper The following abbreviations are used for collec­ (Haenni & Greve 1995). tors: AN: Alf-Jacob Nilsen, BS: BjomA. Sagvol­ den, B0K: Bj0m 0kland, GB & LG: Gudrun MATERIAL WBakkerud & Lita Greve, LOH & OH: Lars Ove Hansen & Oddvar Hanssen, IGK & LG: Ida Greve Practically all the material dealt with in this study Korsnes & Lita Greve, KB: Kai Berggren, KJ: has been sorted out and assembled by LG from Kjell Ame Johanson, JA: Johannes E. Anonby, collections made by herselfand several other col­ JS & LG: John Skartveit & Lita Greve, JS: John lectors (see «Acknowledgements») in different Skartveit, JS & KHL: John Skartveit & Karl H.

65 Haenni & Greve: New records of Norwegian Scatopsidae (Diptera)

Thunes, LG: Lita Greve, LOH & RM: Lars Ove Han­ A Central and North European species, which ap­ sen & Reidar Mehl, LOH: Lars Ove Hansen, LOH pears to be widespread in Fennoscandia (Anders­ & BS: Lars Ove Hansen & BjomA. Sagvolden, OH son 1982). & JB: Oddvar Hanssen & J.I.I.Batvik, OH: Oddvar Hansen, RP: Reidun Pommeresche, SK: Sverre * Thripomorpha bifida (Zilahi-Sebess, 1956) Kobro, TS & OA: T.S9, I spec., leg. B0K. Thripomorpha ha/teratum (Meigen, 1838) This recently described species (Rotheray & Hors­ HES Elverum: Sagtjern (EIS 55), N, shore oflit­ field 1997) was presently known only from few tle lake, 30 June - 1 July 1998, Id', leg. LG. F0 localities in Scotland. The Norwegian material has Sor-Varanger: near Elvenes (EIS 168), 6 Aug. been compared with type material and found to 1996, Id', leg. JS, Sor-Varanger: Neiden, (EIS be conspecific. The presence of this species in 168), N, on Anthriseus sylvestris, 7 Aug. 1996, Norway can thus be ascertained though only fe­ 8eJeJ 399, leg. JS. males are known from this country. New for This species which is widely distributed in Eu­ Fennoscandia and Denmark. rope including Fennoscandia (Andersson 1982, Krivosheina & Haenni 1986) and was recorded in * Ectaetla c/avlpes (Loew, 1846) our first note under the synonym name Rhegmo­ B0 Kongsberg: SansvNesodden: Fagerstrand (EIS 28), LT, edge ofco­ A poorly known European species, previously re­ niferous/deciduous forests and grassland, 15 - 16 corded in the region from Finland (Hackman 1980) Oct.1990, 19, leg. SK, Asker: Bjorkiis (EIS 28), 24 and Sweden (Andersson 1982). Aug. - 10 Oct. 1995, IeJ, leg. LOH & OH. BV Rollag: Veggli (EIS 35), eN, 29 June 1995, Id' 19, leg. BS. Anapausls rectlnervls Duda, 1928 B0 Kongsberg: Sansv

66 Norw. J. Enlomo/. 47, 65-71. 2000

Svevatn area (EIS 31), boggy area, 28 April - 28 35), 13 July 1995, W, leg. BS. May 1997, 2 spec., leg. JS. MRY 0rsta: near The presence of this species in Norway based on Vatnevann (EIS 68), 16 June 1995, W, leg. LG. a old record by Lundstrom (1913) was considered A common cosmopolitan species, often occuring as dubious though possible in our preceding note under anthropogenic conditions (Lastovka & (Haenni & Greve 1995). The finding of recent Haenni 1981). material allows to reinstateA. scutellata in the list ofNorwegian species. This is a very common spe­ * Api/oscatopse bifi/ata (Walker, 1856) cies in deciduous forests in Central Europe (Haenni 1981, Krivosheina & Haenni 1986), reaching north STI Oppdal: Kongsvoll (EIS 79), MT, 900 m, 28 to Southern Scandinavia. July-12Aug.1992, W, 12-19Aug.1992, W I spec., leg. JS. Api/oscatopse subgraci/is Haenni & Greve, 1995 This rather rarely collected species had been re­ STI Oppdal: Kongsvoll (EIS 79), MT, 900 m, 28 corded till now only from the British Isles and July - 12 Aug. 1992, leJ, 12 - 19 Aug. 1992, 2eJeJ, several Central European countries (Krivosheina Spnenbekken (EIS 79), MT, 1300 m, 19 - 22 Aug. & Haenni 1986). Its presence at Kongsvoll is sur­ 1992, W 19, near Sprrenbekken (EIS 79), MT, prising since it is not a particularly boreal species. 1300 m, 19 - 22 Aug. 1992, hundreds of speci­ New for Fennoscandia and Denmark. mens eJeJ and 99, leg. JS. NNV Flakstad: Silsand­ nes (EIS 136), B, 50 m mixed forest, 14 June 1995, Api/oscatopse f/avico//is (Meigen, 1818) 19, leg.TS & OA. AK Nesodden: Fagerstrand (EIS 28), LT, edge of This newly described species appears to be com­ coniferous/deciduous forests and grassland, 3 - 4 mon in Norway. The record from the Lofoten Is­ Sept. 1995, W 299,5 - 6 Sept. 1995,299, 11 - 12 lands is the second one at a low altitude. Unlike Sept. 1995, W 19, 13 - 14 Sept. 1995, 19, leg. most European Scatopsidae, several species of SK. OS Gausdal: Follebu (EIS 54), 12 Aug. 1995, genus Apiloscatopse have an autumnal flight pe­ 5eJeJ, leg. BS. BV Rollag: Veggli (Eis 35), 13 July riod and may present mass occurrences. This is 1995,6eJeJ 1699, leg. BS. apparently also the case for A. subgracilis as can A common European species of Apiloscatopse, be seen from one ofthe above-mentioned records mainly bound to deciduous forests (Krivosheina from Sprrenbekken. & Haenni 1986). A. flavicollis has a southern dis­ tribution in Scandinavia. Efcooke//a a/btfarsis (Zetterstedt, 1850) (= Cooke//a a/btlarsis Zettersted~ * Api/oscatopse f/avocincta (Duda, 1928) BO Kongsberg: Sansvrer (EIS 27), 26 July 1995, STI Oppdal: Kongsvoll (EIS 79), 900 m, on Arch­ W, leg. BS. RY Ha: Ogna (EIS 7), UTM 32V LK angelica, 28 July 1995, 3eJeJ, MT, 900 m, 28 July 138903, MT, 21 June -17 July 1996, W 19, leg. - 12 Aug. 1992, 3eJeJ, leg. JS. FO Sor-Varanger: IGK & LG, 17 July - 21 Aug. 1996, 5eJeJ, 21 Aug. Ytre Lid (EIS 168), 6 Aug. 1996, IeJ, leg. JS. - 28 Sept. 1996, W, leg. LG. A. flavocincta has a wide distribution in Central and The second and third Norwegian records for this North Europe, including Fennoscandia (Sweden and species which is common and widespread in Eu­ Finland) (Andersson 1982, Krivosheina & Haenni rope (Krivosheina & Haenni 1986), but seems to 1986). Its occurrence in Norway was thus expected. be restricted to the southern regions in Scandina­ via. * Api/oscatopse scute//ata (Loew, 1846) AK Nesodden: Fagerstrand (EIS 28), LT, edge of C%bostema infumatum (Haliday, 1833) coniferous/deciduous forests and grassland, 3 - 4 AK L0renskog: Losby (EIS 29), WT, 28 June - 2 Sept. 1995, leJ, leg. SK. BV Rollag: Veggli (EIS Aug. 1991, W, leg. B0K. HOI Kvam: Svevatn

67 Haenni & Greve: New records of Norwegian Scatopsidae (Diptera) area (EIS 31), MT, oligotrophic bog in pine wood­ by JPH has proved that it is a very distinct spe­ land, 29 July - 26 Aug. 1997, 719, leg. JS & LG. cies, that will be redescribed elsewhere. Its distri­ NTI Lierne: Kveskallen (EIS 108), MT, pine for­ bution is still poorly known, but JPH has seen est, 7 - 26 July 1986,4 aa, leg. OH. material from Central European countries, while the above record is the first from Fennoscandia A mainly Northern European species (Haenni & and Denmark. Greve 1995). Only the HO! record represents a new Norwegian locality. * Colobostema triste (Zetterstedt, 1850) Colobostema ntgrtpenne (Meigen, 1830) B0 Drammen: Vnderlia (EIS 28), MT, pine for­ est with clearings, May 1994, W, leg. LOH. BV o Rade: Tasken (EIS 19), MT, 24 June 1995, 19, Rollag: Rollag (EIS 35), CN, 6 Aug. 1993,2 aa, leg. OH & JB. AK Asker: Bjerkas (EIS 28), MT, leg. BS. RY Ha: Ogna (EIS 7) VTM 32V LK 2 July - 24 Aug. 1995, W, leg. LOH & OH, 138903, MT, 14 May- 21 June 1996, W, leg. IGK Lerenskog: Losby (EIS 29), WT, 2-3 years old & LG, 17 July - 21 Aug. 1996, W, leg. LG. forest cut, 27 June - 1 Aug. 1991, 19, leg. B0K, Nesodden: Fagerstrand (EIS 28), LT, edge of co­ A species widely distributed over all Europe (Kri­ niferous/deciduous forests and grassland, 15 - 16 vosheina & Haenni 1986), already recorded from July 1990, W, 17 - 18 July 1990, W, 30 July - 1 Fennoscandia (Sweden and Finland) and Den­ Aug. 1990, la, 1 - 2 Aug. 1990, la, 24 - 25 June mark. C. triste seems to be restricted to southern 1995, W, 14 - 15 July 1995, W, 18 - 19 Aug. part of Norway. 1996, W,27 - 28 July 1997, W, 28 - 29 July 1997, W, 30 - 31 July 1997, W, 4 - 5 Aug. 1997, W,26 Colobostema sp. - 27 Aug. 1997, W, 4 - 5 July 1998, W, 23 - 24 B0 Buskerud: Sigdal, Heimseterasen (EIS 35), F, July 1998,2 aa, 2 - 3 Aug. 1998, 1a, leg. SK. OS Pinus top, 16 June 1998, 19, leg. JS & KHT. Gausdal: Follebu (EIS 54), 12 Aug. 1995,2 aa leg.BS. BV Rollag: Veggli (EIS 35), 13 July 1995, The unique female specimen from Sigdal has pe­ W, leg. BS, Traen saga (EIS 35), MT, Aug. 1994, culiar genital structures, different from any other W 19, leg.BS. AAY Riser: Torskeberg (EIS 11), known species. However, the females of several 21 July 1995, 3aa, leg. SK. VAY Lindesnes: species (including C. obscuritarse recorded here Lillehavn (EIS 1) VTM 32V LK 876305, LT, 19 from B0 Drammen) are still unknown and females July 1999, 1a, leg. KJ. RY Ha: Ogna (EIS 7) VTM without associated males cannot be correctly iden­ 32V LK 138903, MT, 14 May - 21 June 1996, tified at present. 2aa, 21 June - 17 July 1996, W, leg. IGK & LG, 17 July - 21 Aug. 1996, 2aa 19, leg. LG. HOI • Holoplagia bullata (Edwards, 1925) 144 ppSFI Balestrand: Malsnes (EIS 50), MT, 1 ­ o Rade: Tasken (EIS 19) VTM 32V PL 999794, 29 Aug. 1998, 2aa, leg. GB & LG, Sogndal: MT, natural meadow with old trees, 24 June 1995, Lindborg (EIS 50), WT, young pine stand in de­ W 19, leg. OH & JB. ciduous forest, 7 - 21 June 1996, W, leg. JA. A poorly known European species known till now The commonest species of the genus, which ap­ from few localities in British Isles, Central Eu­ pears to be distributed all over the southern part rope and Finland (Cook 1974, Krivosheina & of Norway. Haenni 1986). Probably myrrnecophilous. The Norwegian locality is an untouched meadow * Colobostema obscuritarse (Strobl, 1898) with many large, partly hollow trees (Tilia cO/'data, B0 Drammen: Vnderlia (EIS 28), MT, pine for­ Quercus sp., A/nus g/utinosa) and dead trees as est with clearings, May 1994, W, leg. LOH. well. Many rare beetles have been caught there This species has long been synonymized with C. (Oddvar Hanssen, pers.comm.). triste (Zett.) but the examination of type material

68 Norw. J. Enlomol. 47, 65-11. 2000

* H%p/agia transversa/is (Loew, 1846) * Swammerdame//a acuta Cook, 1956 AK Nesodden: Fagerstrand (EIS 28), LT, edge of RY Ha: Ogna(EIS 7) UTM 32V LK 138903, MT, coniferous/deciduous forests and grassland, 6 - 7 14 May - 21 June 1996,70'0' 5W, leg. IGK & Oct. 1998, 19, leg. SK. BO Kongsberg: Sansva:r LG, 17 July - 21 Aug. 1996,30'0', leg. LG. (EIS 27), 26 July1995, 10', 19, leg. BS. BV Rollag: A Central and North European species, previously Veggli (EIS 35), CN, 19 June 1995, 19, leg. BS, recorded from Sweden and Finland (Krivosheina Veggli (EIS 35), CN, 29 June 1995, 10' 19, leg. & Haenni 1986). Doubts about the identity offe­ BS ( I 9 MHNN). TEI Tinn: Rjukan (EIS 26), males of this species and the closely related S. July 1995, 20'0', leg. BS (10' MHNN). AAI adercotris Cook inclined us to treat them together Bygland: Heddevika (EIS 9), MT, 16 May - 11 as Swamerdamella sp. in our first note (Haenni & June 1997,20'0', leg. KB (10' MHNN). RY Ha: Greve, 1995). Discovery ofmales ofboth species Ogna (EIS 7) UTM 32V LK 138903, MT, 17 July in recently collected material allows us now to - 21 Aug. 1996, 10', leg. LG. ascertain the presence ofboth species in Norway. H. transversalis has a wide European distribution (Krivosheina & Haenni 1986). In the North this * Swammerdame//a adercotris Cook, 1972 mynnecophilous species has been recorded from BO Kongsberg: Sansva:r (EIS 27), 26 July 1995, Finland, Sweden and Denmark (Andersson 1982) and 10', leg. BS. BV Rollag: Veggli (EIS 35), CN, 29 its presence in Norway could therefore be expected. June 1995, 10', leg. BS.

Cobo/dia fuscipes (Meigen, 1830) This poorly known species has been now recorded from several countries in Central and North Euro­ AK Nesodden: Fagerstrand (EIS 28), LT, edge of pe (Krivosheina & Haenni 1986). It is apparently coniferous / deciduous forests and grassland, 10 ­ more generally distributed than mentioned by 11 July 1997, 10', I - 2 Sept. 1997, I 0', leg. SK. Andersson (1982). See discussion under S. acuta. OS Gausdal: Follebu (EIS 54), 12Aug. 1995, 10', leg. BS, Bodal (EIS 54), 12 Aug. 1995,20'0', leg. BS. BO Kongsberg: Sansva:r (EIS 27), 26 July Swammerdame//a sp. (acuta/adercotris) 1995, I 0' 5 W, leg. BS. BV Rollag: Veggli (EIS AK Enebakk: Losby (EIS 29), WT, 10-15 years 35), 13 July 1995,90'0' 2 W, leg. BS, Veggli (EIS old forest cut, 26 June - 30 July 1991, 19, leg. 35), CN, 29 June 1995,30'0' 4 W, leg. BS. TEI B0K. OS Gausdal: Bodal (EIS 54), 12 Aug. 1995, Tinn: Hakanes (EIS 26), MT, Sept. 1995, 10', leg. 19, leg. BS, Follebu (EIS 54), 12 Aug. 1995, 10' BS. RY Finnoy: Sevheim (EIS 14), YT, 26. May­ 299, leg. BS. BO Kongsberg: Sansva:r (EIS 27), 26 June 1995, 10', leg. JS. 26 July 1995, 599, leg. BS, Buskerud: Sigdal, Heimseterasen (EIS 35), F, Pinus top, 17 July A common cosmopolitan species widespread over 1999, 19, leg. JS & KHT. BV Rollag: Veggli (EIS all Europe (Krivosheina & Haenni 1986) in both 35), CN, 19 June 1995, 19, leg. BS, Veggli (EIS anthropogenic and natural environments. JPH has 35), CN, 29 June 1995, 699, leg. BS, Veggli (EIS seen material from practically all European countries. 35),9 July 1995, 19, leg. BS, Viirviken (EIS 35), MT, Aug. 1994,299, leg. LOH & BS. VE Vale: Rhexoza richardsi Freeman, 1985 Langoya (EIS 19), MT, basiphilous pine forest, 2 AK Enebakk: Losby (EIS 29), WT, 10-15 years - 28 May 1991,19, leg. LOH. old forest cut, 26 June - 30 July 1991, 19, leg. The above material could not be identified at spe­ B0K. cies level due to confusion between females ofS. This poorly known species is still known only from acuta and S. adercotris (see note under S. acuta). the British Isles and Norway (Haenni & Greve 1995). Material from the same locality was re­ ported in the preceding note (Haenni & Greve 1995).

69 Haenni & Greve: New records of Norwegian Sca/opsidae (Dip/era)

Swammerdame//a brevicornis (Meigen, 1830) presented in this article and in Haenni & Greve AK Nesodden: Fagerstrand (EIS 28), LT, edge of (1995) has not been collected specificly with the coniferous/deciduous forests and grassland, 1 - 2 purpose ofstudying Scatopsidae. Our view ofthe Aug. 1990, 1Q,2 - 3 Sept. 1997, IQ, leg. SK, Asker: faunistics of the family in Norway and the distri­ Bjorlcis (EIS 28), MT, 2 July - 24 Aug. 1995, 10', bution ofthe different species should thus still be leg. LOH & OH. OS Gausdal: Follebu (EIS 54), 12 regarded as provisional. Aug. 1995,10' IQ, leg. BS, Bodal (EIS 54), 12Aug. Haenni & Greve (1995) recorded 19 species from 1995, 4d'0' 3W,leg. BS. Be Hurum: Toftehohnen Norway, and the authors supposed that the number (EIS 19), MT, dry meadow at seashore, 1 Sept. - 26 of species occuring in this country should be Oct. 1991, IQ, leg. LOH, Drammen: Underlia (EIS around 30 species. In fact, no less than 13 additio­ 28), MT, pine forest with clearings, June 1992, 10', nal species are present in the material dealt with leg. LOH, Kongsberg: Sansvrer (EIS 27), 26 July in this paper, i. e. Ectaetia christii Rotheray & 1995, 4d'0' 699, leg. BS. BV Rollag: Veggli (EIS Horsfie1d, E. clavipes (Loew), E. platyscelis 35), 13 July 1995,499, leg. BS, Veggli (EIS 35), (Loew), Thripomorpha bifida (Zilahi-Sebess), CN, 19 June 1995, 10', leg. BS. TEY Bamble: Apiloscatopse bifilata (Haliday), A. flavocincta Langoya (EIS 11), MT, at seashore, 11 June - 31 (Duda), Apiloscatopse scutellata (Loew), Colo­ July 1995, 30'0', leg. LOH & RM. RY Ha: Ogna bostema obscuritarse (Strobl), C. triste (Zetter­ (EIS 7) UTM 32V LK 138903, MT, 14 May - 21 stedt), Holoplagia bullata (Edwards), H. transver­ June 1996, 10' ?lQ, leg. IGK & LG, Ogna (EIS 7), salis (Loew), Swammerdamella acuta Cook and UTM32VLK 138903,17 July-21 Aug. 1996, 10', S. adercotris Cook, and the total number of spe­ leg. LG. HOY Bomlo: near Olavskolen (EIS 22), cies recorded from Norway is 31 approximately N, 18 Aug. 1992, IQ, leg. LG. as expected. In comparison, 37 species have been A widespread and common species all over Eu­ recorded from Sweden (Andersson 1982) and 26 rope (including Fennoscandia) and Western Palae­ species from Finland (Hackmann 1980). arctic region (Krivosheina & Haenni 1986). It is interesting to note thatmost ofthe newly recor­ ded species are European in general distribution Swammerdame//a genypodis Cook, 1972 and seem to reach their northern limit in southern BV Rollag: Veggli (EIS 35), CN, 29 June 1995, Scandinavia, including the southern parts ofNor­ 2W, leg. BS (1QMHNN). AAI Bygland: Hedde­ way. vika (EIS 9), MT, 1 - 29 July 1998, 10', leg. KB. Seven species (Arthria analis. Thripomorpha hal­ The above records are the 2nd and 3rd Norwegian teratum, Scatopse notata. Apiloscatopse subgra­ records ofthis rare species mentioned for the first cilis, Colobostema infumatum, Coboldiafuscipes, time by Haenni & Greve (1995). All known loca­ Swammerdamella brevicornis) appear to be gen­ lities are located in the southemmost region ofNor­ erally distributed in Norway. Only 5 species (Thri­ way. pomorpha verralli, Rhegmoclemina vaginata, Scatopse lapponica, Apiloscatopse bifilata andA. DISCUSSION flavocincta) are apparently restricted to the north­ ern and central parts ofthe country (North ofTron­ Scatopsidae are minute unconspicuous midges delag), but two ofthem (T. verralli andA. bifilata) usually unfrequently encountered in the field, irres­ are European species whose absence in southern pective ofthe method ofcollecting. Their lifespan Norway probably reflects only insufficient collec­ is short, mass occurences are rare, except in ge­ tion. The remaining nineteen species appear to be nus Apiloscatopse (see under A. subgracilis) and restricted to the southern part of Norway (South in the anthropogenic species S. notata and C. ofTrondelag), though at least two ofthem (Swam­ fuscipes, and to collect a numerous and varied merdamella acuta and S. adercotris) may prob­ material is a large effort. Before discussing the ably also occur in northern part ofthe country. results, it should bear in mind that the material

70 Norw. J. Enlomo/. 47, 65-71. 2000

Among the new records are several poorly known Haenni, J.-P.1981. Contribution ala connaissance de la or rarely collected species: Ectaetia christii, E. faune des Scatopsidae (Diptera) de Suisse. 1. Le platyscelis, Apiloscatopse bifilata, Colobostema genre Apiloscatopse Cook. Mitt. schweiz. ent. Ges. obscuritarse and Holoplagia bullata which all are 54, 257 - 276. Haenni, J.-P. & Greve, L. 1995. Faunistic note about known from few widely separated localities. Both Norwegian Scatopsidae (Diptera), with description species of Ectaetia and H. bullata are very prob­ of a new species. Fauna norv. Ser. B 42, 71-82. ably saproxylic species bound to old trees and may Krivosheina, N. P. & Haenni, J.- P. 1986. Family Scatop­ be indicative of biotopes of conservation value. sidae. Pp. 297 - 310 in S06s, A. & Papp, L. (Eds.), This is also the case for Rhexoza richardsi and Catalogue Palaeartic Diptera. Sciaridae - Anisopo­ Swammerdamella genypodis. didae. Akademiai Kiad6. Budapest. Lastovka, P. & Haenni, J.-P. 1981. Scatopse globuli­ Some additional species can still be expected to cauda sp.n. from the European Alps, with notes on occur in Norway species, particularly in genus S. notata (Diptera, Scatopsidae). Acta ent. Bohe­ Anapausis, of which 5 species are known from moslov. 78,340 - 347. southern Sweden (Andersson 1982) in contrast to Lundstrom, C. 1913. XVI. Diptera Nematocera. Ver­ only one recorded from Norway. Some species zeichnis mehrerer von Dr. E. Strand in Norwegen occurring in marshy habitats in neighbouring Swe­ gesammelten Diptera Nematocera. In: Strand, E. den like Thripomorpha paludicola Enderlein and Neue Beitrage zur Arthropodenfauna Norwegens Ferneiella incompleta (Verrall), andParascatopse nebst gelegentiichen Bemerkungen iiber deutsche Acten. XVI-XX. Nyt Mag. Naturvidenskab. 51,310­ litorea (Edwards), a halophilous species ofthe sea­ 319. shore can also be expected to occur in Norway. Rotheray, G. E. & Horsfield, D. 1997. Ectaetia christii sp. n., a Scottish species similar to Ectaetia clavipes Acknowledgements. We are grateful to Johannes E. (Diptera, Scatopsidae). Dipterists Digest (2nd ser.) Anonby, Leikanger, Kai Berggren, Kristiansand, Lars 4,41- 44. Ove Hansen, Zoological Museum, University of Oslo, Schoyen, WM. 1889. Supplement til H.Siebke·s Enu­ Oddvar Hanssen, NINA, Trondheim, K. A. Johanson, meratio Insectorum Norvegicorum, Fasc.IV (Dipte­ Naturhistoriska Riksmuseet, Stockholm, Sverre Kobro, ra). Christiania Videmskab. Selskab. Forhandl. 12, Norwegian Crop Research Institute, As, Alf-Jacob Nil­ 10. sen, Hidra, BjomA. Sagvolden, Rollag, John Skartveit, Siebke, H. 1877. Enumeratio Insectorum Norvegicorum Zoological Museum, University of Bergen and Bjom Fasciculum IV Catalogum Dipterorum Continen­ 0kland, Norwegian Forest Institute, As has provided tern. 225 pp. A.WBrogger, Christiania. us with material. Some of the material is from of the Zetterstedt, J.W 1838. Sectio Tertia. Diptera. Pp. 477­ research programme «Forest ecology and multiple use», 868 in : Insecta Lapponica. Lipsiae (= Leipzig). and some is from the forest biodiversity project «Miljo­ Zetterstedt, J.W 1850. Diptera Scandinaviae disposita registrering i Skog». We also wish to thank Graham E. et descripta. 9, 3367 - 3710. Ex officina lundberg­ Rotheray, National Museums of Scotland, Edinburgh, iana, Lundae (=Lund). who kindly lent us type material ofEctaetia christi. Received 20 November 1999, References accepted 12 March 2000. Andersson, H. 1978. Revision of the Scatopsidae (Di­ ptera) in Zetterstedfs collection, with lectotype de­ signations. Ent. Scand. 9, 235 - 236. Andersson, H. 1982. De svenska artema av myggfa­ miljema Synneuridae, Canthyloscelidae och Scatop­ sidae. Ent. Tidskr. 103, 5-11. Cook, E.F. 1974. A Synopsis of the Scatopsidae of the Palaearctic. Part Ill. The Scatopsini. J. Nat. Hist. 8, 61 - 100. Hackman, W 1980. A check-list ofthe Finnish Diptera. I. Nematocera and Brachycera s. str. Notulae ent. 60, 17-48.

7 1 Short communication

Malacomyia sciomyzina (Haliday, M. sciomyzina was recorded from Sweden by 1833) (Diptera, Coelopidae) recorded Wahlgren (1917) from Skane and Bohusliin. Ardo from Norway (1957) mentions M. sciomyzina in his lists of Diptera from marine shore dune ecosystem, but he gives no new localities. Lita Greve & Kjell Arne Johanson Gorodkov (1984) placed the genus Malacomyia with the single species M. sciomyzina in the Fam­ Greve, L. & Johanson, K.A. 2000. Malacomyia sciomy­ ily Helcomyzidae together with the genera Helco­ zina (Haliday, 1833) (Diptera, Coelopidae) recorded from Norway. Norw. 1. Entomol. 47, 72. myza Curtis and Heterocheila Rondani. Here we follow McAlpine (1998) who places the genus The coelopid fly Malacomyia sciomyzina (Haliday, Malacomyia together with the genus Coelopa in 1833) (Diptera, Coelopidae) is recorded for the first time the family Coelopidae. from Norway. Two males were caught on light near Lil1ehavn in Lindesnes on 20 August 1999. References

Key words: Malacomyia sciomyzina, Coelopidae, Di­ Ardo, P. 1957. Studies in the marine shore dune eco­ ptera, Norway system with special reference to the dipterous fauna. Opusc. ent. Suppl. XlV, 255 pp. Lita Greve &KjellArne Johanson, University ofBergen, Gorodkov, K.B. 1984. Family Coelopidae. Pp. 151-152 Zoological Institute, Zoological Museum, Museplass 3, in S06s, A. & Papp, L. (Eds), Catalogue ofPalae­ N-5007 Bergen, Norway. arctic Diptera. Yo1.9. Akademiai Kiad6, Budapest. Greve, L. 1999. Litt om tangfluer - Familien Coelopidae. Insekt-Nytt 24 (1), 15-20. The fly Malacomyia sciomyzina (Haliday, 1833) McAlpine, D. 1998. 3.31.Family Coelopidae. Pp. 335­ is recorded for the first time from Norway. Two 340 in Papp, L. & Darvas, B. (Eds.), Contributions to a Manual ofPalaearctic Diptera. Higher Brachy­ males were collected in VAY Lindesnes: Lillehavn cera. Yol.3. Science Herald, Budapest. (EIS 1), 20 August 1999 in a light-trap by K.AJ. Stubbs, A. & Chandler, p.1. 1978. A Dipterist' s Hand­ The locality is open bedrock with patches ofsmall book. The Amateur Entomologist Yol. 15, 255 pp. numbers ofPinus, Calluna and different grass spe­ Wahlgren, E. 1917. Diptera 2. Andra underordningen. cies. The weather was warm and rainy, with strong Flugor Cyclorapha. Andre gruppen Schizophora. onshore wind. Our specimens were collected at Fam. 7. TAngflugor. Coelopidae. Svensk Insektfauna approximately 300 m from the drift line about 10 11,160-163. m a.s.l. They were most probably drifted by wind. Received 15 November 1999, M. sciomyzina looks very much like the species accepted 14 January 2000 in the genus Coelopa, but is separated by the ex­ tensive hairing and the bristle on the anepisternum, the rounded postnotopleural ridge and the bare metasternum, in Coelopa the anepisternum is bare, the postnotopleural ridge is sharply margined and the metasternum is setulose.The distribution ofthe species in the genus Coelopa in Norway is treated by Greve (1999). M. sciomyzina is mentioned as a characteristic species occurring associated with the drift line where seaweed - wrack is the major component. However, the seaweed should be moist and thick (Stubbs & Chandler 1978).

72 Norw. j Enlomo/. 47, 73-75. 2000

New records of spiders (Araneae) from the Oslofjord area, SE Norway

Erling Hauge & Lars Ove Hansen

Hauge, E. & Hansen, L.O. 2000. New records of spiders (Aranea) from the Oslofjord area, SE Norway. Norw. 1. Entomol. 47, 73-75.

The spiders (Araneae) Theridion mystaceum L. Koch, 1870 (Theridiidae) and Linyphia tenuipalpis Simon, 1884 (Linyphiidae) are reported for the first time in Norway. Interesting records of six other species are also presented, including the second Norwegian record ofSyedra gracilis (Menge, 1866) (Linyphiidae). The distribution for each species are briefly discussed.

Key-words: Araneae, Oslofjord, Linyphia tenuipalpis, Theridion mystaceum, Syedra gracilis.

Erling Hauge, Zoological Museum, University ofBergen, Museplass 3, N-5007 Bergen, Norway. Lars Ove Hansen, Zoological Museum, University ofOslo, Sars gate 1, N-0562 Oslo, Norway.

INTRODUCTION THE SPECIES Hauge & Hansen (1991) presented a list of99 spe­ Clubionidae cies of spiders from six small islands in the mid­ Phi/odromlls dispar Walckenaer, 1825 dle ofthe Oslofjord. Several ofthese species cer­ tainly have their northwestern limit of distribu­ A singe CJ was taken at B0 Hurum: Tofteholmen tion at these islands, and surprisingly, five of the (EIS 19), MT. This species is previously known species were previously not reported from Nor­ from coastal areas of SE Norway (Strand 1901, way. The present study gives further records from Hauge & Midtgaard 1986), but an older record these islands, together with an additional record exists from NTI Trondheim (Storm 1898). In Swe­ from the mainland. den it is distributed in coastal areas from Skane to Bohusliin and Uppland, including Gland and Got­ A brief account of the flora and geology of the land (Tullgren 1944). localities are presented by Hansen & Ligaard (1992) and Hauge & Hansen (1991). The EIS-grid Salticidae numbers are given according to 0lcland (1977) and regional abbreviations are according to 0kland Sa/ticlls zebranells (C.L. Koch, 1837) (1981). The second author is responsible for all 1CJ and 599 were taken at B0 Hurum: Tofteholmen the records. The following abbreviations are used (EIS 19), MT, and anotherCJ at VE Vale: Langeya in the text: MT = Malaise-trap operated in the pe­ (EIS 19), MT. The few previous records of this riod 2 May - 26 Oct. 1991, SN = sweep-netting in species in Norway are restricted to the Oslofjord the period 1 July - 12 August 1990. The material area: AK Vestby and VE Tjeme (Hauge 1989). is deposited at the Zoological Museum, Univer­ sity ofBergen. Dendryphantes rlldis (Sundevall, 1833) A single CJ was taken at B0 Hurum: Melen (EIS 19), SN. Only old records of this species are pre­ viously known from Norway:. AK Christiania

73 Hauge & Hansen. New records of spiders (Aranea) from SE Norway omegn, Asker and Eidsvoll, and B0 Drammen Porrhomma microphthalmum (O.P.-Cam­ (Collet 1875). bridge, 1891) A single 9 was taken at B0 Humm: Melen (EIS Agelenidae 19), SN. Recently this species was recorded as new to Norway from Akershus, i.e. AK Ski, Kroer and Agelena labyrinthica (Clerck, 1757) As (Folvik 1994). Palmgren (1975) reported it 2eJeJ and 19 were taken at VE Vale: Lang0j'a (EIS from the S and SW areas of Finland. 19), MT, and 6eJeJ and 19 at Sande: Kommers0j'a (EIS 19), MT. In Norway this species is recorded Syedra gracilis (Menge, 1866) from coastal areas from VAY Kristiansand to AK Oslo, and east to 0 Halden and Hvaler (Hauge A single 9 were taken at B0 Humm: Rarnvikhol­ 1989). The Norwegian records probably represent men (EIS 19) (SN). The species has previously the NW limit of distribution of the species in Eu­ been recorded once from Norway (Waaler 1967), rope. In the Nordic countries it is restricted to S and is seemingly very rare elsewhere in Europe Sweden and the SW coast ofFinland, as well as in (Heimer & Nentwig 1991). In Britain there is a Osterbotten (Kronestedt 1983). This is a continen­ few records north to SE Scotland (Locket et a1. tal species found most frequently in sunny forest 1974). ecotones (Palmgren 1977, Heimer & Nentwig 1991 ). DISCUSSSION Theridiidae A total of 118 species are recorded from these small Theridion mystaceum L. Koch, 1870 islands (Waa1er 1971, Hauge & Hansen 1991, un­ A single eJ was taken at B0 R.0yken: Kinnartangen pub1. records). This represents more than 22% of (EIS 28), MT. This species has previously not been all species ofspiders hitherto recorded in Norway recorded from Norway. It is widely distributed in (Hauge 1989). These islands may be considered Europe (Heimer & Nentwig 1991), and well unique from a national point of wiew, due to a known in E Fennoscandia north to Lappland combination of favourable climate, diverse geo­ (Palmgren 1974). logical origin, and in beeing very little influenced by human activities. This is probably the main rea­ Linyphiidae sons why many species ofspiders and also insects have their northwestern limit of distribution in Linyphia tenuipalpis Simon, 1884 these areas. All these islands are totaly or partly Two eJeJ were taken at B0 Hurum: Tofteholmen protected as nature reserves today. (EIS 19), MT. This is the first Norwegian record of the species. It was reported for the first time in Acknowledgements. We are greatly indebted to lnger Denmark by Toft (1977), but has 01ater been taken Hansen who sewed and repaired some of the Malaise­ both in coastal and interior areas of the country traps and to Geir E.E. Soli and Ketil Aakra for com­ (Toft 1980). The Finnish records are restricted to ments on the manuscript. the S and SW coast including the island ofAland, and eastwards to the Karelian isthmus in Russia References (Palmgren 1977, Lehtinen et a1. 1979). In Sweden Collett, R. 1875. Oversigt afNorgesAraneider.1. Salti­ it is found mostly in coastal areas north to Bohus­ grarue, Citigrarue. Christ. Vidensk.-Selsk. Forhandl. Uin and Sodermanland, also in some distance from 1875, 1-35. the coast (Kronestedt 1983). It is widely distrib­ Folvik, E. 1994. Porrhomma microphtalmum (O.P.­ uted in W Europe south to Portugal, but seems to Cambridge, 1891) (Araneae, Linyphiidae) recorded be scarcer in the NW areas (van Heldingen 1969). new to Norway. Fauna norv. Ser. B 41, 91-92. Hansen, L.O. & Ligaard, S. 1992. Coleoptera from six small islands in the middle Oslofjord, SE Norway.

74 Norw. J. Entomol. 41, 13-15. 2000

Fauna norv. Ser. B. 39, 23-31. rassidae. 138pp. Entomologiska fOreningen, Stock­ Hauge, E. & Hansen, L.O. 1991. Spiders (Araneae) from holm. six small islands in the middle Oslofjord, SE Nor­ Waaler, P.E 1967. A collection of spiders from Son, way. Fauna norv.Ser.B 38, 45-52. Norway. Norsk ent. Tidsskr. 14,91-93. Hauge, E. & Midtgaard, F. 1986. Spiders (Araneae) in Waaler, P.E 1971. Spiders new to Norway. Norsk ent. the Malaise traps from two islands in the Oslofjord, Tidsskr.18, 17-24. Norway. Fauna norv. Ser. B 33, 98-102. Hauge, E. 1989. An annotiated check-list ofNorwegian 4 spiders (Araneae). Insecta Norvegiae 4, 1--40. Received October 1999, Heimer, S. & Nentwig, W 1991. Spinnen Mitteleuropas. accepted 15 March 2000 543 pp. Paul Pareys VerIag, Berlin & Hamburg. Heldingen, PJ. van. 1969. A reclassification ofthe spe­ cies of Linyphia Latreille based on the functioning of the genitalia (Araneida, Linyphiidae). I. Zool. Verh. Leiden 105, 1-303. Kronestedt, T. 1983. Spindlar pi! 01ands Stora alvar. Ent. Tidskr. 104, 183-212. Lehtinen, P.T., Koponen, S. & Saaristo, M. 1979. Stud­ ies on the spider fauna of the southwestern archi­ pelago of Finland n. The Aland mainland and the island ofEckerii. Memo. Soc. Fauna Flora fenn. 55, 33-52. Locket, G.H., Millidge, A.E & Merrett, P. 1974. Brit­ ish Spiders Ill. 314 pp. Ray Society, London. 0ldand, J. 1977. Utt om biogeografiske metoder, og noen nye data om utbredelse av stavtege, Ranatra linearia, og vannskorpion, Nepa cinerea, i Norge. Fauna (Oslo) 30, 145-167. 0k:1and, K.A. 1981. Inndeling av Norge til bruk ved biogeografiske oppgaver - et revidert Strand-sys­ tem. Fauna (Oslo) 34,167-178. Palmgren, P. 1974. Die Spinnenfauna Finnlands und Ostfennoskandiens V. Theridiidae und Nesticidae. Fauna fennica 26, 1-54. Palmgren, P. 1975. Die Spinnenfauna Finnlands und OstfennoskandiensVI. Linyphiidae I. Fauna fenni­ ca 28, 1-102. Palmgren, P. 1977. Die Spinnenfauna Finnlands und Ostfennoskandiens VIII. Fauna fennica 30, 1-50. Storm,Y. 1898. Iagttagelser over Arachnider im siid­ lichen Norwegen. K. norske Vidensk. Selsk. Skr. 1898 (7), 1-10. Strand, E. 1901. Bemerkungen iiber Norwegische Laterigraden nebst Beschreibungen drei neuer oder wenig bekannter Arten. Abh. naturforsch. Ges. Giir­ litz 23, 1-15. loft, S. 1977. Spindlere (Arachnida) fra Anholt. Fau­ nistiske undersfllgelser piAnholt 8. Flora og Fauna 83 (1),19-21. Toft, S. 1980. Linyphia tenuipalpis Simon (Araneae: Linyphiidae) pa dejyske heder. Ent. Meddr. 48,17­ 18. Tullgren, A. 1944. Svensk spindelfauna 3. Fam. 1--4. Salticidae, Ihomisidae, Philodromidae och Euspar-

75 Short communication

The family Signiphoridae The species is a secondary parasite of annoured (Hymenoptera, Chalcidoidea) in scale insects ofthe family Diaspididae (Hemiptera, Norway Coccoidea), probably as an endoparasitoid on other chalcids (Tryapitsyn 1978). Lars Ove Hansen Acknowledgements. I am greatly indebted to Geir E.E. Soli for comments on the manuscript, to Morten Falck for help during the field work, and to 0stensjovannets Hansen, L. O. 2000. The family Signiphoridae (Hyme­ Venner, Oslo, who financed a part of this study. noptera, Chalcidoidea) in Norway. Norw. J. Entomol 47,76. References Fitton, M.G., Graham, M.W.R. de Y., Boucek, Z.R.J., A species ofthe familiy Signiphoridae has been recorded Fergusson, N.D.M., Huddleston, T., Quinland, J. & for the first time in Norway. The minute chalcid wasp Richards, O. W. 1978. A Check List of British In­ Thysanus ater Haliday in Walker, 1840 was recorded in sects, Hymenoptera (2nd. ed.). Handbooks for the Drammen (B0) in 1995 and in Oslo (AK) 1996. Biolo­ identification of British Insects. 11(4), 1-159. gy and distribution ofthe species are briefly discussed. Gauld, I. & Bolton, B. (eds). 1988. The Hymenoptera. British Museum (Natural History). 332 pp. Oxford Key words: Signiphoridae, Chalcidoidea, Thysanus ater, University Press. Norway. Hansson, C. 1991. A Catalogue ofthe Chalcidoidea (Hy­ menoptera) described by C.G. Thomson, with a Lars Ove Hansen, University of Oslo, Zoological Mu­ checklist ofSwedish species. Ent. Scand. Suppl. 38, seum, Sarsgt. 1, N-0562 Oslo, Norway. 1-70. Landin, B.-O. 1971. Hymenoptera Parasitica Chalcidoi­ dea. Pp. 755-895 in Fiiltfauna, Insekter 2 (2), 381­ The family Signiphoridae has been recorded for 1055. Natur och Kultur, Stockholm. the first time in Norway. This family includes Tryapitsyn, Y.A. 1978 (english edition 1988). 17. Family minute chalcid wasps ranging from 0.5 to 2.0 mm Signiphoridae (Thysanidae). Keys to the Insects of in length, with compact or long and flat bodies the European part of the USSR. 3 (2), 513-516. (Tryapitsyn 1978, Gauld & Bolton 1988). The colour is nonnally black, but occasionally with Received 7 March 2000, orange or yellow areas. Wings are fully developed, accepted 23 March 2000 usually with relative long marginal fringes. A single specimen of T. ater was recorded at AK Oslo: 0stensj0vannet, Oppsalskrenten (EIS 28), August 1996 (malaise-trap), leg. Morten Falck & LOH, and another specimen at B0 Drammen: Underlia (EIS 28), July 1995, (malaise-trap), leg. LOH. Both specimens are deposited in the collec­ tions at the Zoological Museum of Oslo. Thysanus ater is recorded from Moldavia, N Cau­ casus, C Asia and W Europe including the British Isles (Tryapitsyn 1978, Fitton et al. 1978). How­ ever, Hansson (1991) did not include the species in his list of Swedish species, but according to Landin (1971), the family is represented by two species in Sweden.

76 Norw. J. Entomo/. 41, 11-88. 2000

New records of spiders (Araneae) from Norway with notes on epigynal characters of Philodromus fuscomarginatus (De Geer) (Philodromidae) and Araneus sturmi (Hahn) (Araneidae)

Kjetil Aakra

Aakra, K. 2000. New records of spiders (Araneae) from Norway with notes on epigynal characters ofPhilodromus fuscomarginatus (De Geer) (Philodromidae) and Araneus sturmi (Hahn) (Araneidae). Norw. 1. Entomol. 47, 77-88.

New records are presented for a number ofspecies collected in western and eastern parts ofsouthern Norway. Saaristoafirma (O.P.-Cambridge, 1905) is new to Fennoscandia. Dipoena torva (Thorell, 1875), Entelecara flavipes (Blackwall, 1834) and Theridion pinastri L. Koch, 1972 are new to Nor­ way. Drassyllus pusillus (c. L. Koch, ,1833), Cercidia prominens (Westring, 1851), Lasaeola tristis (Hahn, 1833), Heliophanus flavipes (Hahn, 1832), Theridion mystaceum L. Koch, 1870 and Sisicus apertus (Holm, 1939) are new to western Norway. The first record ofOedothorax agrestis (Blackwall, 1853) in south Norway and the second record ofDendryphantes rodis (Sundevall, 1832) in over lOO years are reported. Notes on distribution and habitat preferences of the species are given. Variable epigynal characters of Philodromus fuscomarginatus (De Geer, 1778) and Araneus sturmi (Hahn, 1833) are illustrated and briefly discussed.

Key Words: Araneae, Norway.

Kjetil Aakra, Museum ofNatural History and Archaeology, Department ofNatural History, NTNU, N-7491 Trondheim, Norway.

INTRODUCTION nick (1998) except for the linyphiids which fol­ low Tanasevitch (1999). The material is deposited The Norwegian spider fauna is still poorly known in the collections ofthe Museum ofZoology, Uni­ and new species are continually discovered. New versity of Bergen, Norway. records are presented here based on collections by the author on Aslmy near Bergen (Aakra 1998) and elsewhere in western Norway as well as ma­ RESULTS AND DISCUSSION terial taken during a pine canopy fogging project Araneidae in eastern Norway carried out by the forest bio­ diversity project «Miljeregistrering i Skog» ofthe Cercidia prominens (Westring, 1851) Norwegian Forest Research Institute financed by Material: HOY Aslmy (EIS 39), 20 May-3 June the Norwegian Ministry ofAgriculture. Two spe­ 1997, 19, pitfall traps in Sphagnum-bog (leg. K. cies found in the latter collection exhibit epigynal Aakra ). characters that are not in overall agreement with illustrations in the literature. These differences are First record from Western Norway. Previously only illustrated and briefly discussed. a few records from Nordland and Troms (Hauge 1989). There are also unpublished records from Abbreviations ofthe faunal provinces ofNorway inner parts of Hordaland (Hauge pers. comm., follow 0k1and (1981), nomenclature follows Plat- Pommeresche 1999). C. prominens is widespread

77 Aakra: New records of spiders (Araneae) from Norway in Sweden (Jonsson pers. comm.), not recorded sweep netting in low vegetation (leg. K. Aakra). from the Atlantic islands (Ashmole 1979, Bengt­ The genitalia ofsome ofthese specimens (Figure son & Hauge 1979, Agnarsson 1996) and rather lA and lB) correspond for the most part to draw­ uncommon in Britain (Locket & Millidge 1953). ings provided by Wiehle (1931), Tullgren (1952), Hanggi et al. (1995) reported it from several dif­ Locket & Millidge (1953), Levi (1973) and Ro­ ferent situations, including raised bogs, moist mea­ berts (1995). The scapus of the epigyne (follow­ dows and pine forests. ing the terminology of Levi 1971) is coiled as in Figure 1A and the posterior aspect ofthe lamellae Araneus s/urmi (Hahn, 1833) (Figure lB) is very similar to Figure 10 in Levi (1973), but not Tullgren (1952: Figure 2la). The Material: BV Sigdal (EIS 35),19 June 1999, 1 9, median apophysis ofthe male palpal organs fit the 24 June 1999, 1d, 19,25 June 1999,1 d, 26 June description of both Levi and Tullgren. In the fe­ 1999,2 dd, 8. July 1999,399,24 July 1999,19, males two basic variations of the epigyne exists. 25 July 1999, 1 9, 26 July 1999, 1 9, all from There are differences in both the scapus which canopy fogging, crown of pines (leg. 1. Skartveit coils the opposite way (Figure 1C) as often illus­ and K. Thunes). HOY Oster0y: Herlandsnesjane trated for A. triguttatus (Fabricius) and the lamel­ Nature Reserve (EIS 40), 11 June 1999, 299, lae (Figure ID) which here resemble the drawing

Figure 1. Aranel/s stl/rmi. Variation of epigynal characters in some specimens from ventral (A and C) and posterioventral perspective (8 and D).

78 Norw. J. Entomo/. 41, 11-88. 2000 provided by Tullgren (1952) for A. sturmi. tral parts of Sweden (Jonsson pers. comm.) but has not been recorded from Finland (Palmgren The two females from Ostemy appear intermedi­ 1977). In Britain there are scattered records from ate between the two variations noted above. Both southern and central parts (Locket et al. 1974). have the scapus coiled as in a «normal» A. sturmi Otherwise known from continental Europe and (sensu Levi 1973), one has the lamellae of a "nor­ eastwards to Russia and Ukraine (Mikhailov mal" A. sturmi, while the lamellae of the second 1997). Probably reaches its northwestern limit of resemble those of the other females noted above. distribution in SE Norway. Apparently, both types of coiling have been re­ corded in the literature before, although the varia­ Usually associated with bushes and vegetation in tion has not been commented upon. Miller (1971) open sites (Roberts 1987, Maurer & Hanggi 1990), indicated that the coiling in the two species was but evidently also on conifers in northern Europe different while both Wiehle (1931) and Tullgren as the current record suggests. (1952) indicate (albeit vaguely) that the coiling in both species is similar. The illustration of Punda Hypse/istes jacksoni (0. P.-Cambridge, 1902) (1975) show a coiling similar to the one reported here (Figure 1C). The distinguishing external geni­ Material: HOY Aslmy (EIS 39), 21 April-2 June tal features ofA. sturmi and A. triguttatus are there­ 1997, 4 aa, pitfall traps, Sphagnum-bog (leg. K. fore reduced to the shape of the lamellae (see Aakra). Tullgren 1952, and Levi 1973). Second record from Norway. The first was from an eutrophic low-alpine dwarf-shrub heath in Gnaphosidae Jotunheirnen in Oppland, central Norway (Hauge & Refseth 1979). There are also unpublished re­ Drassy//us pusH/us (C. L. Koch, 1833) cords from a bog on Stord (Greve & Hauge 1989) Material: HOY Askey (EIS 39), 12 June-26 June and from inner parts of Hordaland county (Pom­ 1996,2 aa, pitfall traps, Sphagnum-bog (leg. K. meresche 1999). Hjacksoni is generally a north­ Aakra). ern species, known from central and northern parts ofSweden (Jamtland to Tome Lappmark, Jonsson D. pusillus (Figure 2) has previously been recorded pers. comm.), most frequent in northern regions from Norway only once, in eastern parts of the in Finland (Palmgren 1976) and otherwise in nor­ country (Strand 1904). In the 0ygarden archipe­ them parts ofAsia and America (Marusik & Leech lago west ofAslmy it is relatively common (Hauge 1993). The species is also reported from the Brit­ pers. comm.). There are also an unpublished record ish Isles (Roberts 1987), but apparently lacking from Rogaland (Martens 1982). In Sweden it from southern Germany (Blick & Scheidler 1991). ranges north to Uppland (Jonsson pers. comm.), and is rather widespread in Finland (Palmgren H jacksoni is associated with damp, open habi­ 1943). tats (Palmgren 1965, Snazell 1982, Platen et al. 1991) as well as lake shores, birch wood margins, No consensus on the habitat preferences of this wet moss (Palmgren 1976) and Calluna and Erica species seems to have been reached (Grimm 1983). dominated situations (Janssen 1991). On Aslmy It has been found in both moist fields, forests re­ the species was only found in a raised bog domi­ mains and meadows (Miller 1967). nated by Sphagnum and it may be a good charac­ ter species ofpeatlands in coastal parts ofwestern Linyphiidae Norway. Marusik & Leech (1993) did not give any Ente/ecara f/avipes (Blackwall, 1834) detailed information on habitat preferences but mentioned that it could be found in different types Material: BV Sigdal (EIS 35), 19 June 1999, 1a, ofhabitats in both lowlands and mountainous tun­ canopy fogging, crown of pine (leg. J. Skartveit dra. Holm (1983) reported the species as locally & K. Thunes). abundant. New to Norway. The species is known from cen- 79 OB

. D o6ewI 'snll!snd SnjjASS8JO 'l am6!:1

,{OM10N W01/ (BOBU01II) slap/ds /0 splo:Jal MaN :OlJfOll Norw. J. Enlomo! 41, 11-88. 2000

Oedothorax agrestis (Blackwall, 1853) shores. Indeed, the locality in Heriandsnesjane cor­ responds exactly to the habitat descriptions of Material: HOY Osteroy: Heriandsnesjane Nature Palmgren (1976), Hauge (1976) and Maurer & Reserve (EIS 40),13 August 1999,2 crcr, 2 QQ, I Hiinggi (1990). juvenile, manually collected from stony lakeshore. 0. agrestis is distinguished from other Oedothorax First record from southern Norway since Strand species in the region by the large ventral apophy­ (1902) who reported it from Suldal in inner Roga­ sis on the palpal tibia (Figure 3A) and by the un­ land. There is also one record from Finnmark modified carapace in the male (Figure 3B) while (Hauge 1976). The species is widespread in Swe­ the female (epigyne in Figure 3C) is distinguished den (Jonsson pers. comm.), Finland (Palmgren by internal genitalia (see Roberts 1987) or by as­ 1976), Ireland and Great Britain (Locket et al. sociation with males. A peculiar anomaly was 1974), continental parts of Europe (Maurer & noted regarding the male palps: the smallest tibial Hiinggi 1990) and ranges as far east as South-Si­ apophysis is reduced in length, but only on the beria (Esyunin & Efimik 1996). left pedipalp (Figures 3D and 3E). The species appears to be common on stony lake­

c

/

E B

~i~ure 3. Oedothorax agrestis. cJ palpus (A) and carapace (8) and epigyne (C). Left (D) and right (E) tibial apohyses of both cJ specimens examined.

8 1 Aakra: New records of spiders (Araneae) from Norway

Saaristoa firma (O.P.-Cambridge, 1905) Most records are from moist moss-dominated (pre­ dominantly Sphagnum) situations in various types Material: HOY Aslmy (EIS 39), 22 August-21 offorests (Holm 1968, Kritscher 1971/72, Casemir September 1997; 2 aa, 1 9, pitfall traps, Alnusl 1976, Hauge 1977). On Aslmy S. corniger was Pinus forest. Bf3mlo (EIS 22), 3 August 1998, 19, found in various habitats with lush vegetation, manually collected in deciduous forest, (leg. E. suggesting that it is not strictly sphagnicol as ear­ Hauge). lier records may indicate. Other authors (Casemir First record from Norway, indeed S. firma is new 1976, von Broen 1985, Hiinggi et al. 1995) have to Fennoscandia as it has not been reported from reached the same conclusion. Sweden (Jonsson and Kronestedt pers. comm.), nor from Finland (Koponen pers. comm.). There Sisicus apertus (Holm, 1939) are a few unpublished records from Denmark (Toft pers comm.). In central Europe the species seems Material: HOY Aslmy (EIS 39), 2 September-23 to be most common in the mountainous regions April 1997, 4 aa, pitfall traps, AlnuslPinus forest including the Alps, sometimes being locally abun­ (leg. K. Aakra). dant (Thaler 1981). In northern Europe records First record from western parts ofNorway, previ­ tend to be more infrequent and collections sites ously only known from SE areas of the country more scattered (Locket et al. 1974). It is not known (Hauge 1989), reaching Tome lappmark in Swe­ from Russia or Siberia (Tanasevitch pers. comm.). den (Jonsson pers. comm.) and known from most Records from a private database (kindly made parts ofFinland (Palmgren 1975). It has not been available by T. Blick) indicate that this species may recorded from Great Britain or Ireland (Merrett et be found in a range of different situations, often al. 1985, Merrett 1995) and is rare in high-alti­ in Sphagnum-dominated sites, different humid fo­ tude areas in continental Europe (Thaler 1969, rests types (predominantly spruce and mixed spru­ 1993). Widespread throughout northern Asia to ce forests) and caves. Casemir (1976) took S. firma Alaska, the Rocky Mountains and the North-Ame­ in moist moss in spruce forests while Kropf & rican Atlantic coast (Eskov 1994). Horak (1996) suggested that litter of deciduous In Norway the species has previously been taken forests constitutes the principal habitat ofthe spe­ from pine forests, mixed pine forests, TUccinium­ CIes. Pinetum mixed forest, a bog (Hauge & Wiger 1980, Tveit & Hauge 1983) and forest-fire areas Sintula corniger (Blackwall, 1856) (Hauge & Kvamme 1983). It has also been found in margins of mires and in Sphagnum in spruce Material: HOY Aslmy (EIS 39), 17 April-22 Sep­ forests (Palmgren 1975), moss-dominated conif­ tember 1996,9 May-23 May 1997, 5 aa, 299, erous forests (Thaler 1993) and Hylocomium-moss pitfall traps, various habitats (leg. K. Aakra). in a birch forest (Holm 1939). Second official record ofthis species from south­ ern Norway, previously known from Tysfjord and Philodromidae Narvik in north-eastern Nordland (Hauge 1989). It is also known from Stord (Greve & Hauge 1989) Phi/odromus fuscomarginatus (De Geer 1778) and inner parts ofHordaland county (Pommeres­ Material: BV Sigdal (EIS 35): 5 June 1998, I 9, che 1999). In Sweden the species has been found hand collecting, beneath bark ofdead pine, 10 June from Uppland to Lycksele Lappmark (Jonsson 1998, 2 aa, canopy fogging, crown of pine, 24 pers. comm.), in Finland north to central parts of June 1999, la, canopy fogging, crown of pine the country (Palmgren 1975). Apparently widely (leg. 1. Skartveit and K. Thunes). distributed in most parts ofEurope from the north The female illustrated here (Figure 4) appears to to the south, but rarely reported (Kritscher 1971/ be conspecific with Philodromusfuscomarginatus 72), although it may be locally abundant (Casemir sensu Tullgren (1944), Punda (1975) and Heimer 1976).

82 Norw. J Entomol. 41, 11-88. 2000

& Nentwig (1991) but differs in some respects in Salticidae epigynal characters. The posterior lateral borders Dendryphantes rudis (Sundevall, 1832) of the epigyne (Figure 4) are closer together in my specimen than indicated by Tullgren (1944) Material: BV Sigdal (EIS 35), 5 June 1998, 1 9, and Heimer & Nentwig (1991) and is more in ac­ canopy fogging, crown of pine (leg. J. Skartveit cordance with the drawing of Punda. The central and K. Thunes). ridge (<

He/iophanus f/avipes (Hahn, 1832) Material: HOI Etne: Ljusnes (EIS 23),8 July 1999, 1 (J, handpicked from grazed pasture (leg. K. Aakra). New to western Norway. Previously known from the Oslofjord area (Hauge 1989, Andersen & Hauge 1995). Occurs north to Medelpad in Swe­ den (Jonsson pers. comm.), rather common in southern and central parts of Finland (Palmgren 1943) and widespread in Britain and Ireland (Locket et a1. 1974) and the rest of the Palearctic region (Maurer & Hanggi 1990, Esyunin & Efimik 1996). H. flavipes is usually associated with open grassy fields with varying degrees of moisture (Maurer & Hiinggi 1990), a thermophilous species accord­ ing to Braun (1969) and Buchar (1975).

Figure 4. Phi/odromus fuscomarginatus. Epigyne. Hairs omitted from epigyne for clarity, except for the stout hairs on the central ridge.

83 Aakra: New records of spiders (Araneae) from Norway

Theridiidae (1995). This is probably the result of different interpretations by the various authors or variations Dipoena lorva (Thorell, 1875) in the level of sclerotisation. Material: BV Sigdal (EIS 35), 26 June 1998, 1 9, canopy fogging, trunk ofold pine (leg. J. Skartveit Lasaeo/a lrislis (Hahn, 1833) and K. Thunes). Material: BOY Ostemy (EIS 40): Herlandsnesja­ First record from Norway. D. torva is known from ne Nature Reserve, 11 June and 12 August 1999, Si:idermanland, Ostergi:itland and Uppsala in Swe­ 3 dd, sweep netting from low vegetation on bogs den (Jonsson pers. comm.), southern Finland and wetlands (leg. K. Aakra) (Palmgren 1974), northern Great Britain (Roberts 1995) and central Europe and is known as far east First record from western Norway, previously as Middle-Siberia (Esyunin & Efimik 1996, Mik­ known from the Oslofjord area (Hauge 1989, Hau­ hailov 1997), but rare throughout its known range. ge & Hansen 1991,2000). Recorded north to Norr­ botten in Sweden (Jonsson pers. comm.) and ap­ The species is restricted to older trees, predomi­ proximately the same latitude in Finland (Palm­ nantly pines in northern Europe, where is spins a gren 1974). In Great Britain and Ireland it is quite small web in deep fissures in the bark which is local and restricted to southern counties (Locket used to catch wood ants (Bratton 1991, Roberts et al. 1974). L. tristis is otherwise known from 1995, Platenpers. comm.). Presence ofolder trees continental Europe and Russia (Platnick 1998). with deeply structured bark covering and wood The current record probably represent the north­ ants would thus be necessary for this species (Ro­ westemmost limit of distribution of this species. berts 1995). In consequence, there can be little doubt that modern forestry practices represent a It is most common in Calluna and low vegetation threat to this species (Palmgren 1974). This is sub­ although it can also be found on young conifers stantiated by the fact that the Norwegian speci­ (Palmgren 1974). men was taken on a 250 year old pine (measured at breast height) measuring 17.5 m in height and Theridion myslaceum L. Koch, 1870 having a 8.5 m canopy diameter (Skartveit pers. Material: BOY Ask0y (EIS 39), 9 June 1996, 19, comm.). 15 June 1996,2 dd and 2 99,20 July 1996, 1 The shape of the dark markings above the central subadult d, 19,31 October 1996, 299, all manu­ fissure ofthe epigyne (Figure 5) differs somewhat ally collected from road-cuts and vertical rock from the illustrations in Miller (1967) and Roberts sides. Bergen: Kalfaret (EIS 39), 5 May 1999, 1 d, 1 subadult d and 3 juveniles, manual collect­ ing from road-cut. B0mlo, near Olavsskolen (EIS 22), 13 August 1996, 1 9, manual collecting in road cut (leg. K. Aakra). New to Western Norway, the only other known record is from the Oslofjord region (Hauge & Han­ sen 2000). Available evidence suggests that T. mystaceum has often been confused with the closely related spe­ cies T. melanurum Hahn, at least in Sweden (Jons­ son 1995). No T. melanurum specimens were de­ posited in the collections of the Zoological Mu­ seum in Bergen. Given the commonness of T. mys­ taceum in western parts of Norway (from where the single report of T. melanurum originated, see Figure 5. Dipoena lorva. Epigyne.

84 Norw. J. Entomo/. 41, 77-88. 2000

Hauge 1989) the misidentification has most prob­ way. The current Norwegian records indicate that ably occurred in Norway as well. T. melanu11lm T. mystaceum is very common in road-cuts and should therefore be omitted from the Norwegian similar natural habitats. check-list until identifyable specimens are col­ The distinguishing features of the epigynes of T. lected. mystaceum and T. melanu11lm are illustrated in T. mystaceum is usually found on and near build­ Figure 6. The ducts of T. mystaceum are characte­ ings, T. melanu11lm preferring somewhat drier situ­ rised by the more rounded epigynal socket in ations, although this does not seem to hold for which the seminal ducts may be visible. These Britain (Parker 1990). Locket et a1. (1974) indica­ ducts are also more extensively coiled in T. mysta­ ted that T. melanu11lm could also be found amongst ceum than in T. melanu11lm. Detailed examina­ stones on the Continent, while T. mystaceum is tion is needed to distinguish the males (Locket et most commonly found on tree trunks, evergreen a1. 1974, Roberts 1985). bushes and occasionally on the outside of houses and sheds. These habitat descriptions are of little Thendlon pinastri L. Koch, 1872 value in separating the species according to Ro­ berts (1985). In Britain T. mystaceum is the com­ Material: BV Sigdal (EIS 35), 24 June 1999, 1 cr, monest of the two, T. melanu11lm being less fre­ canopy fogging, crown of pine (leg. 1. Skartveit quent in the north which is consistent with the & K. Thunes.). apparent absence ofthis species from western Nor- New to Norway. The species occurs north to Upp-

B

c

Figure 6. The epigyne and vulvae of T. mystaceum (A and C) and T. me/anurum (B and D). Drawings of T. me/anurum after Roberts (1985).

85 Aakra: New records of spiders (Araneae) from Norway land in Sweden (Jonsson pers. comm.) and is Bratton, J. H. 1991 (ed.). British Red Data Books. 3. apparently rather common in southern Finland Invertebrates other than insects. 253 pp. J.N.C.C., (Palmgren 1974), but was only recently discov­ Peterborough.. ered in Great Britain (Murphy & Murphy 1979) Braun, R. 1969. Zur Autoko10gie und Phanologie der Spinnen (Araneida) des Naturschutzgebietes where it is rare (Bratton 1991). The species is also «Mainzer Sand». G1eichzeitig ein Beitrag zur known from southern and central Europe and KenntnisderThermophilie bei Spinnen. Mainzer na­ throughout Asia to Japan (Esyunin & Efimik turw. Arch. 8, 193 - 289. 1996). Broen, von, B. 1985. Bemerkungen iiber einige Nach­ weise selten gefundener Spinnearten (Araneae). Although it has been found in semi-open habitats Zoo1. Anz., Jena 214, 151 - 156. with Calluna and Juniperus (Palmgren 1974, Buchar, 1. 1975. Arachnofauna Bohmens und ihr Murphy & Murphy 1979), coniferous forests ap­ thermophi1er Bestandtei1. Vest. cs1. Spo1. zoo1. 39, pear to be preferred (Palmgren 1974, Maurer & 241 - 250. Hanggi 1990). Casemir, H. 1976. Beitrag zur Hocmoor-Spinnenfauna des Hohen Venns (Hautes Fagnes) zwischen Nord­ Acknowledgements. Special thanks to Theo Blick for eife1 und Ardennen. Decheniana 129, 38 - 72. much valuable help regarding the ecology and distri­ Collett, R. 1875. Oversigt over Norges Araneida 1. Forh. bution of Saaristoa firma, to Lars J. Jonsson for pro­ VidenskSe1sk. Krist. 1875,225 - 259. viding information on the distribution ofthe species in Donda1e, C. D. & Redner, 1. H. 1975. Thefuscomargi­ Sweden, to John Skartveit and Karl Thunes ofthe for­ natus and histrio groups ofthe spider genus Philo­ est biodiversity project «Miljoregistrering i Skog» for dromus in North America (Araneida: Thomisidae). providing access to their spider material, to Havard Can. Ent. 107,369 - 384. Bjordal, Fylkesmannen i Horda1and, Mi1jovemavdelin­ Eskov, K. Y. 1994. Catalogue of the linyphiid spiders ga, for permission to publish in advance the material of northern Asia. 144 pp. Pensoft publishers Sofia from the Nature Reserve at Herda1snesjane, to Erling - Moscow. Hauge and Reidun Pommeresche for their valuable Esyunin, S. L. & Efimik, V. E. 1996. Catalogue of the comments and information on various species and lastly spiders (Arachnida, Aranei) of the Ura1s. 229 pp. to Torbjorn Kronestedt, Soren Toft, Andrei Tanasevitch Ed. K. G. Mikahi10v. Moscow: KMK Scientifc and Seppo Koponen for kindly providing information Press Ltd. 1996. on the occurence (or absence) of S. firma in their re­ Greve, L & Hauge, E. 1989. Insekt- og edderkopp­ spective countries. faunaen pa myrer i Hordaland. 35 pp. Rapport terrestrisk oko10gi nr. 49. Zoological Museum, Uni­ versity of Bergen, December 1989. REFERENCES Grimm, U. 1983. Revision der Gnaphosidae Mitte1euro­ Aakra, K. 1998. Epigeic spiders ofthe island ofAskoy, pas. 265 pp. Dissertation des Doktorgrades des Fac­ western Norway. 148 pp. Cand. ScientThesis, Zoo­ bereichs Bio10gie der Universitet Hamburg. Ham­ logical Museum, University of Bergen, June 1998. burg 1983. Agnarsson,1. 1996. Is1enskar kongu1rer. 175 pp. Fjo1rit Hanggi, A., Stockli, E. & Nentwig, W. 1995. Lebens­ Natturufrredistofnunar 31. raume Mitte1europaischer Spinnen. Charactherisi­ Andersen, T. & Hauge, E. 1995. Pitfall catches of spi­ erung der Labensraume der hauFiguresten Spinnen­ ders (Araneae) from proposed nature reserves on arten Mitte1europas und der mit diesen vergesell­ Tjome, Vestfo1d, SE Norway. Fauna norv. Ser. B schaften Arten. Miscellanea Faunistica He1vetiae 42,1 - 10. 4, Neuchate1. Centre suisse de cartographie de la Ashmo1e, N. P. 1979. The spider fauna of Shetland and faune. 460 pp. its zoogeographic context. Proc. Royal Soc. Edin­ Hauge, E. 1976. Araneae from Finnmark, Norway. burgh 78B, 63 - 122. Norw. 1. Entomo1. 23, 121 - 125. Bengtson, S.-A. & Hauge, E. 1979. Terrestrial inverte­ Hauge, E. 1977. The spider fauna oftwo forest habitats brates of the Faroe Islands: 1. Spiders (Araneae): in northern Norway. Astarte 10,93 - 101. Check-list, distribution, and habitats. Fauna norv. Hauge, E. 1989. An annotated check-list ofNorwegian Ser. B 26, 59 - 83. spiders (Araneae). Insecta norvegia 4, 1 - 40. Blick, T. & Scheid1er, M. 1991. Kommentierte Arten­ Hauge, E. & Hansen, L. O. 1991. Spiders (Araneae) liste der Spinnen Bayerens (Araneae). Arachnol. from six small islands in the middle Oslofjord, SE Mitt. 1,27 - 80. Norway. Fauna Norv. Ser. B 38, 45 - 52.

86 Narw J. Enlama/. 47, TT-88. 2000

Hauge, E. & Hansen, L.O. 2000. New records of spi­ Hypselistes, including two new species from Sibe­ ders (Aranea) from the Oslofjord area, SE Norway. ria and the Russian Far East (Araneida: Erigonidae). Norw. J. Entomol. 47, 73 - 75. Can. Ent. 125, 1115 - 1126. Hauge, E. & Kvamme, T. 1983. Spiders from forest­ Maurer, R. & Hanggi, A. 1990. Katalog der Schwei­ fire areas in southeast Norway. Fauna norv. Ser. B zerischen Spinnen. Documenta Faunistica Helvetica 30,39 - 45. 12. Hauge, E. & Refseth, D. 1979. The spider fauna of 5 Merrett, P., Locket, G. H. & Millidge, A. F. 1985. A alpine and subalpine habitats in the Jotuheimen area, check list of British spiders. Bull. Brit. arachnol. southern Norway. Fauna norv. Ser. B 26, 84 - 90. Soc. 6, 381 - 403. Hauge, E. & Wiger, R 1980. The spider fauna (Araneae) Merrett, P. 1995. Eighteen hindred new county records from 12 habitats in the Vassfaretregion, south-east­ of British spiders. Bull. Br. arachnol. Soc. 10, 15 ­ ern Norway. Fauna norv. Ser. B 27, 60 - 67. 18. Heimer, S. & Nentwig, W. 1991. Spinnen Mitteleuropas. Mikhailov, K. G. 1997. Catalogue of the spiders of the Ein Bestimmungsbuch. 543 pp. Verlag Paul Parey. territories of the former Soviet Union (Arachnida, Holm, A. 1939. Neue Spinnen aus Sweden. Beschreib­ Aranei). 416 pp. Moscow: Zoological Museum of ung neuer arten der Familien Drassidae, Theridi­ the Moscow State University. 1997. idae, Linyphiidae und Micryphantidae. Ark. ZooI. Miller, F. 1967. Studien iiber die Kopulationsorgane 31A, I- 38. der Spinnengattung Zelotes, Micaria, Robertus und Holm, A. 1968. A contribution to the spider fauna of Dipoena nebst Beschreibung einiger neuen oder un­ Sweden. Zool. Bidr. Uppsala 37, 183 - 209. vollkommen bekannten Spinnenarten. Acta sc. nat. Holm, A. 1983. Niigot om spindelfaunan i Peljekaise Brno. 1,251- 298. nationalpark. Ent. Tidskr. 104, 13 - 22. Miller, F. 1971. Pavouci - Araneida. Klic zvireny CSSR Hutha, V. & Raatikainen, M. 1974. Spider communiti­ 4,51 - 306. es ofleys and winter cereal fields in Finland. Ann. Murphy 1. & Murphy F. 1979. Theridion pinastri L. Zool. Fennici 11, 97 - 104. Koch, recently found in Britain. Bull. Br. arachnol. Janssen, M. 1991. Spinnen van de Mechelse hiede. Na­ Soc. 4, 314 - 315. tuurhistorisch Maandblad 80, 12 - 16. 0kland, K. 1981. Inndeling av Norge til bruk ved bio­ Jonsson L. 1. 1995. On the spider species of Theridi­ geografiske oppgaver - et revidert Strand system. idae in Sweden (Abstract), p. 223 in. Ruzicka, V. Fauna 34, 167 - 178. th (ed.), Proc. 15 • Colloq. Arachnol. Institue of En­ Palmgren, P. 1943. Die Spinnenfauna Finnlands. 11. tomology, Czech republic 1995. Pisauridae, Oxyopidae, Salticidae, Clubionidae, Kritscher, E. 1971/72. Oreonetides abnormis (Black­ Anyphaenidae, Sparassidae, Ctenidae, Drassidae. wall) 1841 und Sintula corniger (Blackwall) 1856, Acta. ZooI. Fenn. 36, I- 112. (Aran., Linyphiidae), zwei fur Osterreich neue Spin­ Palmgren, P. 1950. Die Spinnenfauna Finnlands und nenarten. Verh. Zool-Botan. Gess., Wien 110/111, Ostfennoskandiens Ill. Xysticidae und Philodromi­ 89 - 94. dae. Acta. Zool. Fenn. 62, 1 - 43. Kropf, C. & Horak, P. 1996. Die Spinnen der Steiermark Palmgren, P. 1965. Die Spinnenfauna der Gegend von (Arachnida, Araneae). Mitt. naturwiss. Ver. Steier­ Kilpisjiirvi in Lappland. Acta Zool. Fenn. 110, I­ mark. Sonderheft, 5 - 112. 70. Levi, H. W. 1971. The diadematus group of the orb­ Palmgren, 1974. Die Spinnenfauna Finnlands und Ost­ weaver genus Araneus north of Mexico (Araneae: fennoskandiens. V. Theridiidae und Nesticidae. Araneidae). Bull. Mus. Comp. Zool. 141, 131 - 179. Fauna Fennica 26, 1 - 54. Levi, H. W. 1973. Small orb-weawers ofthe genus Ara­ Palmgren, P. 1975. Die Spinnenfauna Finnlands und neus north of Mexico (Araneae: Araneidae). Bull. Ostfennoskandiens. VI. Linyphiidae I Fauna Fen­ Mus. Comp. Zool. 145,473- 552. nica 28, I- 102. Locket, G. H. & Millidge, A. F. 1953. British spiders. Palmgren, P. 1976. Die Spinnenfauna Finnlands und 449 pp. Vol. 11. Ray Society, London. Ostfennoskandiens. VII. Linyphiidae 2. Fauna Fen­ Locket, G. H., Millidge, A. F. & P. Merret. 1974. Brit­ nica29,1-126. ish Spiders Ill. 314 pp. London, Ray Society. Palmgren, P. 1977. Die Spinnenfauna Finnlands und Martens, T. 1982. Registrering av landlevende everte­ Ostfennoskandiens VlII. Argyronetidae, Agelen­ brater pii Kiirste og Ogney i Rogaland. 89 pp. Zoo­ diae, Hahniidae, Dictynidae, Amaurobiidae, Tita­ logical Museum, University ofBergen. Rapport Nr. noecidae, Segestridae, Pholcidae And Sicariidae. 45. Fauna Fennica 30, I- 50. Marusik, Y. M. & Leechr, R. 1993. The spider genus Parker, J. R. 1990. The lifecycle of Theridion mysta­

87 Aakra: New records of spiders (Araneae) from Norway

ceum L. Koch. Newslt. Br. arachno!. Soc. 58,2 - 3. ansand and Setesdalen, S. Norway. Fauna norv. Ser. Platen, R. , Moritz, M. & von Broen, B. 1991. Liste der B 31,23 - 45. Webspinnen- und Weberknechten (Arach.: Aranei­ Wiehle, H. 1931. Spinnentiere oder Arachnoidea. VI: da, Opilionida) des Berliner Raumes und ihre Aus­ Agelenidae - Araneidae. 136 pp. In: Dahl, F. Die wertung fiir naturschutzzwecke (Rote liste). Land­ Tierwelts Deutschlands. schaftentwicklung und Umweltsforschung S 6, 169 - 205. Platnick, N. 1. 1998. Advances in Spider Taxonomy Received 12 January 2000, 1992 - 1995. With redescriptions 1940 - 1980. 976 accepted 30 March 2000 pp. New York Entomological Society 1997. Pommeresche, R. 1999. Diversitet, sarnrnfunnstrukturer og habitatspreferanser hos epigeiske edderkopper i ulike vegetasjonstyper innen Geitaknottane natur­ reservat, indre Hordaland. 70 pp. Cand. scient. the­ sis. Zoological Museum, University of Bergen, spring 1999. Punda, H. 1975. Pajaki borow sosnowychi. 91 pp. Polska Akademia Nauk. Instytut Zoologi. Roberts, M. 1. 1985. The spiders of Great Britain and Ireland. 229 pp. Harley Books, Colchester. Roberts, M. 1. 1987. The spiders of Great Britain and Ireland Vo!. 2. Linyphiidae and checklist. 204 pp. Harley Books, Colchester. England. Roberts, M. J. 1995. Spiders of Britain & Northern Europe. 383 pp. Collins Field Guide. HarperCol­ linsPublishers. Snazell, R. 1982. Habitat preferences of some spiders on heathland in southern England. Bull. Br. arachn. Soc. 5, 352 - 360. Strand, E. 1902. Theridiiden aus dem westlichen Nor­ wegen. Bergens Mus. Arb. 6, I - 23. Strand, E. 1904. Die Dictyniden, Dysderiden, Drassi­ den, Clubioniden, undAgeleniden der Collet'schen Spinnensarnlung. Forh. VidenskSelsk Krist. 1904, I- 16. Tanasevitch, A. V. 1999. Linyphiid spiders ofthe world. http://www.andtan.newmail.ru/list/linyphiidae.htrn Thaler, K. 1969. Ober einge wenig bekannte Zwegspin­ nen aus Tirol (Arachn., Araneae. Erigonidae). Ber. nat. -med. Ver. Innsbruck 57,195 - 219. Thaler, K. 1981. Ober Oreonetides quadridentatus (Wunderlich (1972) novo comb. (Arachnida: Aranei, Linyphiidae). Arch. Sc. Geneve 34,143 - 152. Thaler, K. 1993. Ober wenig bekannte Zwergspinnen aus den Alpen - IX (Arachnida: Aranei, Linyphii­ dae: Erigoninae). Rev. suisse Zool. 100,641 - 654. Tullgren, A. 1944. Salticidae, Thomisidae, Philodro­ midae och Eusparrassidae. 138 pp. Svensk Spinde1­ fauna 3, Fam. I - 4. Stcokholm (Entomologiske f6reningen). Tullgren, A. 1952. Zur Kenntnis schwedischer Spinnen. Ent. Tidskr. 73, 151 - 177. Tveit, L. & Hauge, E. 1983. The spider fauna ofKristi­

88 Norw. J. Enlomo! 41, 89-93. 2000

New records of Norwegian Lauxaniidae (Diptera)

Uta Greve

Greve, L. 2000. New records of Norwegian Lauxaniidae (Diptera). Norw. 1. Entomo1.47, 89-93.

Seven species of lauxaniid flies (Diptera, Lauxaniidae) Homoneura tenera (Loew, 1846), Aulogas­ tromyia anisodactyla (Loew, 1845), Lyciella decipiens (Loew, 1847), Lyciella platycephala (Loew, 1847), Lyciella subpallidiventris Papp, 1978, Lyciella vittata (Walker, 1849) and Sapromyza quadricincta Becker, 1895 are recorded for the first time from Norway. Lyciella subpallidiventris and Sapromyza quadricincta are both recorded new to Fennoscandia and Denmark. Remarks are made on the distribution of Tricholauxania praeusta (Fal1en, 1820) in Norway.

Uta Greve, Zoological Museum, Zoological Institute. University ofBergen, Museplass 3, NO-5007 Bergen. Norway.

INTRODUCTION (Scopoli, 1763). Thus only nineteen species ofthe originally twentytwo in Siebke's check list truely The fly family Lauxaniidae (Diptera) is still un­ belong to the family Lauxaniidae. sufficiently known from Norway. Siebke (1877) recorded twentytwo lauxanid species from Nor­ Due to variability in characters used in descrip­ way, not as Lauxaniidae proper, however, fifteen tions in the past, there are many species or spe­ species were listed as genus Sapromyza Fallen and cies-groups that needs revision. A few genera have seven as the genus Lauxania Fabricius. been revised like Minettia by Shatalkin (1998). One should also bear in mind that while males can Number thirteen in the genus Sapromyza listed by have very distinctive characters in the genitalia, Siebke was Sapromyza jlava Lin. (= Linneus, the females often do not, and females of several 1758). This name is today valid as Chyromyajlava species are difficult to determine with certainty. (L.) (Chyromyidae), see Andersson (1971) and is listed as this species by S06s (1984). As number New species from Norway have in the later years fourteen Siebke listed Sapromyza bipunctella been recorded by Rognes (1995), Greve (1997, Zetterstedt (= Zetterstedt, 1847). Andersson (1971) 1999), Greve & Skartveit (1998) and Greve, Pom­ after examination ofZetterstedfs material, decided meresche & Skartveit (1998). The systematic fol­ that S. bipunctella (Zetterstedt, 1847) is a syno­ lows Papp & Shatalkin (1998) and Papp (1984). nym ofanother species in the family Chyromyidae Abbreviations used in the text are: LT= Light trap, viz. Chyromyiaftmorella (Fallen, 1820). As num­ MT= Malaise trap, CT= Collision trap, YT= Yel­ ber fifteen Siebke listed Sapromyza chrysophthal­ low tray. ma Zetterstedt (= Zetterstedt, 1847). Andersson List of abbreviations: AF: Arne Fjellberg, BS: (1971) examined the typematerial of this species Bjorn A. Sagvolden, B0K: Bjorn 0kland, IG: Ida which were all females. Andersson designated a Greve Korsnes, GB: Gudrun W Bakkerud, LG: lectotype from Zetterstedt's material and Anders­ Lita Greve, GAH: Godtfred Anker Halvorsen, son also states that this lectotype belongs to the LOH: Lars Ove Hansen, OH: Oddvar Hanssen, same species as described by Fallen as Sapromyza DWJ: D.WJohansen, JK: Jostein Korsnes, SK: lutea Fallen, 1820. Sapromyza lutea (Fallen, 1820) Sverre Kobro, FM: Fred Midtgaard, AFO: Arne is today considered a synonym of a third species Foldvik, HB: Harald Breilid, HS: Harald Solberg, in the family Chyromyidae Chyromyia oppidana JS: John Skartveit, LS: Liv Sognes, TA: Trond

89 Greve: New records of Norwegian Lallxaniidae (Dip/era)

Andersen. perate deciduous forest, open grassland and a gar­ den (Kobro 1991), and this lighttrap has been run Regional abbreviations are given in accordance throughout every summer since 1984. Lauxaniid with 0kland (1981). flies have, however, only been sorted out for some of the latest years. Subfamily Homoneurinae The light trap was opened in June every year, and Homoneura tenera (Loew, 1846) the first specimens ofA. anisodactyla appeared in Record: TEI Tinn: Hilkanes, Hilkanesodden (EIS early August. Specimens have been caught in the 26), MT, July 1995, Id' 19, leg.BS. months between August and October which sug­ The Malaise trap was operating the whole sum­ gest a flight period in the autumn in southern Nor­ mer, but only two specimens were collected. The way. trap was situated on an open area which earlier A. anisodactyla was described from Germany. It was a small farm, and the forest around consisted has been recorded from Sweden and Finland and of partly conifers and partly deciduous trees. is distributed in most of Europe including Italy In the subfamily Homoneurinae the small black according to Papp (l984a).The male has the outer spines along the costa reach apex of vein R 4+5 claw of the tarsus of the third pair of legs very while in all genera in the subfam. Lauxaniinae enlarged and can be easily recognized because of these spines diminish in size and stop between this character. The female can be recognized on apices of R2+3 and R4+5. H. tenera is a light­ the segment seven ofthe abdomen which is formed yellow coloured species with dark spots on the like a tube and somewhat enlarged. The colour of wings, and on account of the spotted wings it su­ head and body is light yellow. The tip of the an­ perficially looks like Lyciella decempunctata (Fal­ tenna is darkened. len) in the subfamily Lauxaniinae which is a very A. anisodactyla is the only species in this genus in common species in the lowlands ofSouthern Nor­ NW Europe. Shatalkin (1993) described another way. species A. rohdendorfi from Turkmenistan. H. tenera is the first species of the genus Homo­ neura and of the subfamily Homoneurinae recor­ Lycie//a decipiens (Loew, 1847) ded from Norway. The genus Homoneura is the Record: AK Nesodden: Fagerstrand (EIS 28), LT, only genus ofthis subfamily in Fennoscandia and 9 - 10 July 1997, 19, leg. SK. Denmark and four species have been recorded For description of the locality see above. 1. deci­ from Finland (HackmanI980) and three from piens is a yellow species with a shining thorax and Sweden (H. Andersson pers. comm.). H. tenera, yellow antennae and black palpi. The eight ster­ however, is not recorded from Sweden. num of the female abdomen is large and of a dis­ tinctive form see Remm & Elberg (1979). Subfamily Lauxaninae 1. decipiens is recorded from Sweden (H. An­ Au/ogastromyia anisodacty/a (Loew, 1845) dersson pers. comm.), Finland (Hackman, 1980) Records: AK Nesodden: Fagerstrand (EIS 28), LT, and from Denmark (as 1. descipiens (Loew)) by 11-120ct.1994,299,2-3Aug.1995,19,3-4 Reddersen, 1995. Sept. 1995,10,4 - 5 Sept. 1995, 19,24 - 25 Sept. 1995, 1 9, 8 - 9 Oct. 1995, 19, 3 - 4 Aug. 1998, 1 Lycie//a p/atycepha/a ( Loew, 1847) 9,20 - 21 Aug. 1998, Id' 19, leg. SK. Records: AK Nesodden: Fagerstrand (EIS 28), LT, Two males and 8 females ofA. anisodactyla were 4 - 5 Sept.1995, 10, leg. SK. sorted out from material collected in a light trap For description of the locality Nesodden: Fager­ at Akershus: Fagerstrand. The trap was situated in strand, see above. an edge habitat between old coniferous forest, tem­

90 Norw. J. Enlomo/. 47, 89-93. 2000

The females of L. p/atycepha/a have large, ever­ 1986, W 19, leg. FM. MRI Norddal: Fjora (EIS sible organs in the membrane at each side of the 77), MT, 23 June - 18 July 1993, 1 a, 23 June - 18 abdomen. The hind margin of the forth terga of July 1993, W, leg. OH. MRY Hareid: Habakken the abdomen have very long, strong bristles espe­ (EIS 76), MT, 19 - 24 July 1991, 1 9, leg. LG, cially on the side. The males have small outer geni­ Hareid: Krakholen (EIS 75), MT, 1 - 20 May 1990, talia, see Remm & Elberg (1979). 1 9, 20 May - 10 June 1990, 1a, 10 June - 16 July 1990, 4 4 99, leg. JK. NTI Stjordal: Vikan L. p/atycepha/a is recorded from Sweden (H. An­ aa (EIS 92), MT, 25 May - 28 June 1990, 1 9, leg. dersson, pers.comm.) and Finland (Hackman, AFO. 1980). L. vittata has frons and thorax coloured with dis­ Lyciella subpallidiventris Papp, 1978 tinct, longitudinal and alternating stripes in grey Record: AK Nesodden: Fagerstrand (EIS 28), LT, and chocolate brown. The genitalia of both sexes 28 - 29 Aug. 1997, 1 a, leg. SK. are distinctive, see Remm & Elberg (1979). Papp (1978b) redescribed L. vittata, and he also decided For description of the locality see above. L. sub­ that L. quadrivittata (Loew, 1861) (=Sapromyza pallidiventris is very similar to L. pallidiventris quadrivittata Loew, 1861) is a synonym ofL. vit­ (Loew, 1847) and differs in characters ofthe male tata (Walker, 1849). genitalia only. It was described together with L. sty/ata Papp, 1978 from Hungary and Germany All records are from lowlands in southern Nor­ (Papp, 1978). The females ofL. pallidiventris, L. way where L. vittata seems to have a wide distribu­ tion. At Hyllestad, Botnen two MT were operat­ sty/ata and L. subpallidiventris can not be sepa­ rated at present. Dr.Laszlo Papp, Budapest has ing from 25 May untill 14 September 1999, and verified the determination. 36 specimens were collected from May to July. One trap was situated in deciduous forest, the other L. subpallidiventris has been recorded from Hun­ in groups of Cory/us avellana L. Specimens were gary, Germany, Estonia and the British Isles viz. caught between 21 May and 25 July and none be­ England, Ireland/Eire and Scotland (Godfrey tween 25 July and 14 September when the Ma­ 1994), and it is here recorded new from Fenno­ laise trap was taken down. At Hareid, Krakholen scandia and Denmark. one MT operating from early May untill Septem­ ber, 10 specimens were caught between 1 May and Lyciella vittata (Walker, 1849) 16 July, and none later. Records: AK Enebakk: Nordre Boler (EIS 29), According to Papp (1984a) L. vittata has been re­ MT, June 1996, 1 9, leg. HB & HS. B0 Nedre corded from both Finland and Sweden. Eiker: Mjondalen, Miletjern (EIS 28), LT, Ultimo June 1988, 19, leg. DWJ. RY Ha: Ogna (EIS 3), Sapromyzosoma quadricincta (Seeker, 1895) 21 May 1988, 1 9, leg. LG, Ogna (EIS 3), MT, 19 Synonym: Sapromyza bipunctata Meigen, 1830 July - 21 Aug. 1996, 1 9, leg. IG & LG. BOY Bergen (Fana): Flesland (EIS 30), 30 May 1966, Record: B0 Drammen: Underlia (EIS 28), MT, 1 a, leg. AF, Bergen (Asane): Eidsvag, Vollane June 1994,1 a, leg. LOH. (EIS 39), MT, 21 June - 10 July 1986, 19, leg. LG, The male ofthis species has large genitalia «curved det. Hugo Andersson, Os: Gassand, Raudlien (EIS caliper-like towards each other» according to Col­ 30),6 - 28 June 1990, 19, leg. GAH. SFY Gulen: lin (1948). The male from Underlia was compared Brekkestranda (EIS 49) 30 June 1983, 1 9, leg. with material in the Hope collection, Oxford. LG, Hyllestad: Botnen 32 UTM 995917 (EIS 48), MT, 21 June - 25 July 1999,299, 32 UTM 996917, Papp (1984a) reports the distribution to be: Great MT, 21 May - 21 June, 12aa 1999,21 June - 25 Britain, Germany, France, Austria, Hungary, the July, 3 99, leg. LS & LG, Naustdal: Naustdal (EIS former Czechoslovakia, Romania, Bulgaria and 58), MT, 28 May - 3 July 1986, la, 3 - 28 July the former Jugoslavia, and does not mention Fen­ noscandia or Denmark. Thus, S. quadricincta is 9 1 Greve: New records of Norwegian Lauxanlidae (Dip/era) probably new to this area. AK Oslo: Bekke1aget 19 June 1850 1 9 and MRY Gjernnes: Gjernnes (EIS 84)14 Aug. 1843 19. Tricho/auxania praeusta (Fallen,1820) Much new material has been collected which clear­ New records: AKNesodden: Fagerstrand(EIS 28), ly shows that T praeusta is one ofthe more com­ LT, 24 - 25 June 1995, 1 0,5 - 6 Aug. 1995, 1 9, mon species oflauxaniids in the lowlands ofsouth­ 15-16Aug. 1995,10,2-3Aug. 1996,3003W ern Norway. The localities Gjernnes: Gjernnes and 12 - 13 Aug. 1996, 10,29 - 30 Aug. 1996, 1 0, 10 Nordda1: Fjora in the province More and Romsdal - 11 Aug. 1997, 1 9, 18 - 19 Aug. 1997, 10,20­ are the northernmost records. '" 21 Aug. 1997,200 I 9,28 - 29 Aug. 1997 19, T praeusta is known from Sweden (H.Andersson LT, leg. SK, Asker: Bjorkas (EIS 28), MT, 24 Aug. pers.comm.) and Finland (Hackman, 1980). - 10 Octo 1995, 1 0 1 9, leg. LOH & OH. BV Rollag: Traen saga (EIS 35), MT, Aug. 1994, 1 9, leg. BS. VE Tjome: Moutrnarka (EIS 19),22 June Acknowledgements. I thank the following persons who have in various ways helped me collecting material: 1995,2 W1eg. JS. RY Ha: Ogna 32 UTa VLK Trond Andersen, Bergen, Harald Breilid, Bergen, Arne 138903 (EIS 3), MT, 17 July - 21 Aug. 1996,2 Fjellberg, Tjome, Arne Foldvik, Stavanger, Lars Ove 00,2 MT, leg. IG & LG, FiIlllflY: Kvitevik (EIS Hansen, Oslo, Oddvar Hanssen, Trondheim, Godtfred 14), MT, 26 June - 5 August 1995, 1 0 12 W, MT, Anker Halvorsen, Bergen, Ida Greve Korsnes, Bergen, leg. JS. HOY Bergen: Munkebotten (EIS 39), 14 Jostein Korsnes, Kvamsoy, Fred Midtgaard, As, BjOffi July 1987,10, leg. LG, Straume (EIS 30), 31 Aug. A. Sagvolden, Rollag, John Skartveit, Bergen, Liv Sog­ 1979,102 99 leg. TA, (Asane) Vollane, YT, (EIS nes, Hyllestad, Bomen, Harald Solberg, Nannestad and 39),7 - 9 Sept. 1992, 1 9, leg. LG, Bom10: Ka1v­ BjOffi 0kland, As. A special thank to Sverre Kobro, As for his light trapping collection done throughout many nesset (EIS 22), 17 - 18 Aug. 1992, 1 0, leg. LG, years at Fagerstrand: Nesodden, a locality with a rich Fusa: Djupvik (EIS 31), 10 July 1995,200, leg. and very interesting fauna and for placing his material GB & LG. HOI Kvam: Gravdal, Svevatn (EIS for my disposition. I am very grateful to Hugo Anders­ 31), MT, 28 May - 1 July 1997, 32 00 20 W, 1 ­ son, Lund, Sweden for information on the Swedish dis­ 29 July 1997,600599,29 July - 11 Sept.1997, tribution of some species and also to Laszlo Papp, Bu­ 18009 W, 11 Sept.- 29 Oct.l997, Ir:J 19, leg. JS. dapest, Hungary who verified the determination of the SFYHyllestad: Botnen, 32V KN 995917/996917 L. subpallidiventris male. (EIS 48), 2 MT, 21 May - 21 June 1999, 1899,21 June - 25 July 1999,400799,25 July - 7 Sept. References 1999,100019 W, 7 - 14 Sept. 1999, 1 07 W, Andersson, H. 1971. The Swedish species of Chyro­ leg. LS & LG; Naustda1: Naustda1 (EIS 58), MT, myidae (Diptera) with lectotype designations. Ent. 3 - 28 July 1986,19, leg. FM. MRINorddal: Fjora Tidsskr. 92, 95 - 99. (EIS 77), MT ,23 June - 18 July 19931019,18 Godfrey, A. 1994. Lyciella sty/ata Papp and Lyciella July - 11 Sept.1993, 2 Wleg. OH. subpallidiventris Papp (Diptera: Lauxaniidae) new to Britain. Br. 1. Ent. Nat. Hist. 7, 81 - 84. T praeusta was published from 0stfold: Fredriks­ Greve, L.1997. Cnemacantha muscaria (Fallen, 1823) hald, Akershus: Oslo and Enebakk and More og (Dipt.,Lauxaniidae), a species new to Norway. Fau­ Romsdal: Gjernnes by Siebke (1877) as Sapromy­ na norv. Ser. B 44, 78. za prceusta Fallen. This material was mis1abeled Greve, L.1999. Lyciella sty/ata Papp, 1978 (Diptera, in the collection in the Zoological Museum, Uni­ Lauxaniidae) new to Fennoscandia; Lyciella pallidi­ versity ofOslo, and therefore overlooked by Greve ventris (Fallen,1820) and Sapromyza basalis Zet­ terstedt, 1847 (Diptera, Lauxaniidae) new to Nor­ & Skartveit who (1998) published T praeusta as way. Dipterists Digest 6, 33 - 34. a new species for the Norwegian fauna from inner Greve, L. & Skartveit, 1. 1998. Three species ofLauxa­ and outer Rogaland, and inner More og Romsdal. niidae (Diptera) new to Norway and a note on the However, later the following material of T prae­ distribution of Pachycerina seticornis (Fallen, 1820). Fauna norv. Ser. B 45, 110 - 112. usta collected by Siebke were discovered in the Greve, L., Pommeresche, R. & Skartveit, 1. 1998. Pa­ collection in Oslo: 0 Halden: Fredrikshald I 0,

92 Norw. J. Entomol. 47, 89-93. 2000

chycerina pulchra (Loew, 1850) (Diptera, Lauxa­ niidae), a species new to Fennoscandia. Dipterists Digest 5, 95. Hackman, W. 1980. A check list of Finnish Diptera. 11. Cyclorrhapha. Not. ent. 60, 117 - 162. Kobro, S. 1991. Annual variation in abundance ofphoto­ tactic Lepidoptera as indicated by light-trap catches. Fauna norv. Ser. B. 38, I - 4. 0kland, K.A. 1981. Inndeling av Norge til bruk ved biogeografiske oppgaver - et revidert Strand-sys­ tem. Fauna (Oslo) 34, 167 - 178. Papp, L. 1978. Contribution to the revision ofthe Palae­ arctic Lauxaniidae (Diptera). Ann. Hist.-nat. Mus. Nat. Hung. 70,213 - 231. Papp, L. 1984a. Family Lauxaniidae. Pp.193 - 217, in S06s, A. & Papp, L.(eds): Catalogue ofPalaearctic Diptera. Vo!.9, Akademiai Kiad6, Budapest. Papp, L. 1984b. Lauxaniidae (Diptera), newpalaearctic species and taxonomical notes. Acta Zoo!. Hungr. 30,159-177. Papp, L. & Shatalkin, A.!. 1998. Family Lauxaniidae Pp. 383 - 400, in Papp, L. & Darvas, B. (eds): Con­ tributions to a Manual ofPalaeartic Diptera. Vo!. 3, Science Herald, Budapest. Reddersen,1. 1994. Distribution and abundance oflau­ xaniid flies in Danish cereal fields in relation to pes­ ticides, crop and field boundary. Ent. Meddr. 62, 117-128. Remm, E. & Elberg, K. 1979. Terminalia of the Lauxaniidae (Diptera) found in Estonia, Latvia and Lithuania. Dipteroloogilisi Uurimusi, Eesti NSV Tead.Akad., 66 - 117. Rognes, K. 1995. Recent records ofrare flies from Nor­ way (Diptera: Lauxaniidae, Fannidae, Tachinidae). Fauna norv. Ser. B 42, 136 - 137. Siebke, H. 1877. Enumeratio Insectorurn Norvegicorurn Fasciculurn IV. Catalogum Dipterorurn Continen­ tern. 255 pp. A.W. Bmgger, Christiania. Shatalkin, A.I. 1993. New species of Lauxaniidae (Di­ ptera). Russian Entomo!. 1. 2 (3-4), 105 - 118. Shatalkin, AJ. 1998. Palaearctic species of the genus Minettia (Diptera, Lauxaniidae). Zoo!. Jour. 77(7), 809 - 817. S06s, A. 1984. Family Chyromyidae. Pp. 65 - 60, in SOGS, A. & Papp, L. (Eds): Catalogue ofPalaearctic Diptera. Vo!.1 0, Akaderniai Kiad6, Budapest.

i Received 19 January 2000, accepted 20 March 2000

/

93

Norw. J. Entomol. 41. 95-99. 2000

Agyneta mossica (Schikora, 1993) (Araneae, Linyphiidae) in Norway

Kjetil Aakra

Aakra, K. 2000. Agyneta mossica (Schikora, 1993) (Araneae, Linyphiidae) in Norway. Norw. 1. Entomol. 47, 95-99.

All material of Agyneta saxatilis (B1ackwall, 1844) in the collections of the Zoological Museum, Bergen, has been revised and specimens of the recently described and closely allied A. mossica (Schikora, 1993) identified. Illustrations of distinguishing morphological characters and notes on the distribution and habitat preferences of both species are presented. Both species are common within their known area of distribution, A. saxatilis apparently being restricted to southern Norway while A. mossica has a wide distribution having been found from Finnmark to western parts of S Norway.

Key words: Araneae, Agyneta mossica, A. saxatilis.

Kjetil Aakra, Museum ofNatural History andArchaeology, Department ofNatural History, NTNU, N-7491 Trondheim, Norway.

INTRODUCTION Norwegian spiders (Aakra & Hauge in prep.) it became necessary to revised the aformentioned Schikora (1993) described Agyneta (Meioneta) material in order to assess the status ofA. mosscia mossica from Swedish, German and British ma­ in Norway. terial showing minute but distinct differences from its sibling species A. saxatilis (Blackwall, 1844). The two species are also ecologically separated, METODS AND MATERIAL A. mossica being restricted to bogs whereas A. The material examined includes all specimens la­ saxatilis is found in a variety ofhabitats. Further belled «Meioneta saxatilis» (Blackwall) in the examinations of intra- and interpopulational mor­ collection ofthe Zoological Museum, a few addi­ phological variation in both species showed that tional specimens collected by the author in west­ although intergrading specimens did exist the spe­ ern Norway and material kindly provided by O. cies could be reliably separated by the morpho­ Hanssen and F. 0degaard from Vard"" Finnmark. logical differences (Schikora 1995). Schikora Males are differentiated by palpa1 characters and (1995) also elaborated on differences in habitat by differences in cheliceral shape, according to preferences and reportedA. mossica from Norway Schikora (1993, 1995). The presence ofonly a sin­ for the first time (35 km south ofNarvik andVarde, gle paracymbial tooth in A. mossica (Figure I) Finnmark). The material ofA. saxatilis in the col­ seems to be a reliable distinguishing character. lection of the Zoological Museum, University of Females can only be separated by detailed exami­ Bergen, was not examined by Schikora, however, nation of epigynal characters, specifically by the and the distribution ofA. mossica in Norway was degree of curvature of the copulatory ducts (Fig­ therefore uncertain at best. Furthermore, Norwe­ ure 2, also see Schikora 1995). gian records of A. saxatilis was not included in the distribution map of this species (Schikora Schikora originally described the species in the 1995). During work with the red list proposal for genus Meioneta Hull, 1920 but I have chosen to

95 Aakra: Agyneta mossica (Araneae, Linyphiidae) in Norway follow Saaristo (1973) and Russian authors (e.g. April-June 1989, 1a, bog (Greve & hauge 1989). Tanasevitch 1999) who have synonymised Meio­ Ostemy: Herlandsnesjane Nature Reserve (EIS neta with Agyneta Hull, 1911, even if this is not 40),11 June-12. August 1999, 1 <;:>, bog (Aakra & followed in recent catalogues (P1atnick 1998). Djursvoll2000). MRY Haram: L"vs"ya (EIS 76), Abbreviations of fauna1 provinces follow 0k1and 1-18 June 1992, 1 <;:>, bog. NSI Sa1tfjellet: Bj"llii­ (1981). Unless otherwise stated all specimens were da1en (EIS 124), 1-22 July 1974,2 aa, 3 99, open, caught in pitfall traps. sloping ombotrophic bogs, (Hauge 1976); RY Tysvrer: Kiirst" (EIS 14),8-20 May 1981,2 aa, 1 <;:>, bog (Martens 1982).

RESULTS Hauge (1976) reported 2 aa and 6 99 of Meio­ neta saxatilis collected at Bj"lliida1en, Nord1and. Agyneta mossica In the material only 2 aa and 3 99 could be found FN Yard,,: Komagvrer: Hollamyra (EIS 178), 5­ which originated from this site (Hauge pers. 10 July 1998,400, 1 <;:>, pa1sa bog (leg. O. Hanssen comm.). & F. 0degaard). BOY Lindiis: Fonnes (EIS 39), 3-6 June 1978, 1<;:>, bog. Stord: Ig1atj"dn (EIS 23), Agyneta saxatilis AK Frogn: Hii"ya (EIS 28), 3-16 June 1984, 1 <;:>, malaise trap, mixed deciduous forest (Hauge & Midtgaard 1986). Brerum: Ost"ya (EIS 28), ma­ laise trap 30 May-lO June 1984, 1 a, 1 <;:>, transi­ tion sone between bog and forest (Hauge & Midt­ gaard 1986). BOY Os: Hattvik (EIS 31), 8 June 1971, 1 a, 1 <;:>, sieving of moss, mixed forest.

Figure 1. Agyneta mossica (A) and A. saxatllis (B). Figure 2. Agyneta mossica (A) and A. saxatllis (B). Paracymbium of male pal ps. Vulval structures.

96 Norw. J. Entomol. 47, 95-99. 2000

Ostemy, Lono (EIS 40), 4 May-12 June 1973, 1 widespread in Norway (Figure 3), probably oc­ cr, 11-21 July 1973, 1 9, light traps, damp site curring throughout the country. Most records are (Andersen et al. 1980). Os: Bj0rnasen (EIS 31), from coastal parts of western Norway (the most 12 August 1975,19, meadow, 28. May 1975, 19, intensively investigated area in Norway), the other sieving in meadow. B0mlo: Andal (EIS 22), 24 records are from Saltfjellet, Narvik in Northern May-18 September 1977, 1cr. Bergen: Asane (EIS Norway (Schikora 1995) and Finnmark. Itis found 39), 26 June 1980, 19. 0ygarden: Alvheim (EIS throughout Sweden and Finland (Schikora 1995). 39), 21 June - 27 July 1982, 2 crcr, 1 9, gently A. saxatilis on the other hand appears to be a more sloping grassy field. 0ygarden: Hjelme (EIS 39), southern species, in Sweden found north to Upp­ 21 June-8 July 1982,2 crcr, 19, overgrown mea­ land and restricted to southern parts of Finland dow. 0ygarden: Sture (EIS 39), 21 June-8 July (Schikora 1995). Known distribution in Norway 1982, 1 cr, deciduous forest, 8 July-21 July 1982, is at present restricted to coastal areas north to 1 cr, 19, grassy meadow. Os, St. (?) (EIS 31), 27 Hordaland (Figure 4), but any conclusions on the June-ll July 1991, 1cr. Ask0Y: Skansen (EIS 39), northern limit are premature. Schikora (1995) sug­ 29. May - 12. June 1996, 2 crcr, raised bog, 26 gested that this limit is represented by the north­ June-10 July 1996, 4 crcr, 19, young spruce forest ernmost records in Sweden and Finland. (Aakra 1998); HOI Rosendal: Veavag (EIS 31), 10 July 1943, 19. Kvinnherad: JEnesdalen (EIS In regard to habitat choice A. saxatilis is very eury­ 31), 11 May-26 September 1977,5 crcr, 19, mixed topic, Norwegian records span from Sphagnum­ deciduous forest. Voss (EIS 41), 26. June 1978, 1 bogs to transition sones between forests and open cr, forest. Kvam: Geitaknottane Nature Reserve biotopes to various types of forests. According to (EIS 31),12 May-21 September 1997,2 crcr, de­ Schikora ecotones, ruderal sites and meadows ciduous forest; (Pommeresche 1999-not exam­ seem to be preferred. A. mossica on the other hand ined). RI Halandsdalen: Erfjord (EIS 14),26 April­ is consistently found only in bogs or fens and 27 September 1977,4 crcr, 1 9, mixed deciduous should be considered a sphagnophilous species forest. RY Mostemy (EIS 14),5 July-25 Septem­ characteristic of these habitats. ber 1977,6 crcr, 6 99, coniferous forest. Fister, 26 A. mossica is red listed in Germany being threat­ April-27 September 1977, I cr. Tysvrer: Karst0 ened due to habitat loss (Platen et al. 1996). The (EIS 14), 8-30 May 1981, I grased pasture cr, species is not threatened in Norway given our rela­ (Martens 1982). Klepp: Bole (EIS 7), 8 June 1990, tively large quantities of Sphagnum-bogs, but its 1 cr, dune scrub (Folvik 1992). STI Trondheim: stenotopy makes it sensitive and vulnerable to Steinan (EIS 92), 20 July 1975, i moist meadow cr, habitat deterioration, especially in regard to hy­ (leg. B. Larsson). VE Borre: Borrevatn (EIS 19), drology. It would thus be an excellent indicator­ 6 June-20 September 1979, 1cr, moist spruce for­ species of the quality of peatland habitats. est. Brunlanes: Anvik (EIS 19), 7 June-20 Sep­ tember 1979, 1 cr, deciduous (Alnus) forest. Acknowledgements. Thanks to Erling Hauge for pro­ Holmestrand: Eidsfoss (EIS 19), 6 June-20 Sep­ viding habitat descriptions and localities on unpublished tember 1979,7 crcr, 299. VAY Kristiansand: Ytre material and for comments on the manuscript. Also Strandi (EIS 2), 25 June-25 September 1980, 3 thanks to O. Hanssen and F. 0degaard for letting me crcr, 2 99, meadow (Tveit & Hauge 1983). look at their interesting spider material from Vard0.

REFERENCES DISCUSSION Aakra, K. 1998. Epigeic spiders of the island ofAslmy, It is not surprising to find representatives ofAgy­ western Norway. 148 pp. Cand. Scient Thesis, Zoo­ neta mossica in the material examined. For in­ logical Museum, University of Bergen. stance, all records of A. saxatilis from the Polar Aakra, K. & Djursvoll, P. 2000. Terrestre Arthropoder Urals actually referred to A. mossica (Koponen et fra Herlandsnesjane Naturreservat pa Oster0Y, Hor­ daland. Rapport. Fylkesmannen i Hordaland, Milj0­ al. 1997). Our records indicate that A. mossica is

97 86

'ABMJON 'papnl3xa (966~) BJO~!IPS JO pJ03aJ a41 'ABMJON U! UO!lnq!JIS!P UMOU)l 'sf/flexes el8uJIJv 'iI aJn6!.:1 U! UO!lnq!JIS!p UMOU)l 'e:JfssoUJ el8uJIJv '£ aJn6!.:1

;.{eMJON VI (8ep/.llId,.{V/7 '8e8VeJV) e:>!ssow ej8uAfh/ :eJJfev Norw J. Enlomol. 41, 95-99. 2000

vernavdelingen. em and central Europe (Araneae: Linyphiidae). Bull. Andersen, T., Bakke, B. & Klausen, F. E. 1980. Spiders Br. Arachnol. Soc. 10, 65 - 74. '. I, (Araneae) from five damp localities on Osteroy, Tanasevitch, A. V. 1999. Linyphiid spiders ofthe world. Western Norway. Fauna norv. Ser. B 27, 53 - 59. http://www.andtan.newmail.ru/list/linyphiidae.htrn Folvik, E. 1992. The spider fauna ofcoastal sand dunes Tveit, L. & Hauge, E. 1983. The spider fauna ofKristi­ • in southwestern Norway. 65 pp. Cand. Scient. The­ ansand and Setesdalen, S. Norway. Fauna norv. Ser. J sis. Museum of Zoology, University of Bergen, B31,23-45. , Norway. Greve, L & Hauge, E. 1989. Insekt- og edderkopp­ Il faunaen pa myrer i Hordaland. Rapport terrestrisk Received 12 January 2000, .. accepted 30 March 2000 ...., okologi m. 49. 35 pp. Zoological Museum, Univer­ , sity of Bergen. IF. Hauge, E. 1976. Notes on eight species of spiders (Ara­ ~ neae) from the Saltfjellet area, Nordland. Norw. 1. Entomol. 23, 45 - 46. Hauge, E. & Midtgaard, F. 1986. Spiders (Araneae) in Malaise traps from two islands in the Oslofjord, Nor­ way. Fauna norv. Ser. B 33, 98 - 102. Koponen, S., Marusik. Y. M & Tanasevitch, A. V. 1997. New data on the spider fauna ofthe Polar Urals (Ara­ nei). Arthropoda Selecta 6, 109 - 119. Martens, T. 1982. Registrering av landlevende everte­ brater pa Karsto og Ognoy i Rogaland. Rapport Nr. 45. 89 pp. Laboratorium for ferskvannsokologi og innlandsfiske, Zoological Museum, University of Bergen. 0k1and, K. 1981. Inndeling av Norge til bruk ved bio­ geogratiske oppgaver - et revidert Strand system. Fauna 34, 167 - 178. Platen, R., Blick, T., Malten, A. & Sacher, P. (unter Mitarbeit zahlreicher Fachleute) 1996. Rote Liste der Webspinnen Deutschlands (Arachnida: Ara­ neae). Arachnol. Mitt. ll, 5-31 Basel. Platnick, N. I. 1998. Advances in spider taxonomy 1992 - 1995. With redescriptions 1940 - 1980. 976 pp. Ent. Soc. Arner. Mus Nat. Hist., New York. Pomrneresche, R. 1999. Diversitet, sanunfunnstrukturer og habitatspreferanser hos epigeiske edderkopper i ulike vegetasjonstyper innen Geitaknottane natur­ reservat, indre Hordaland. 70 pp. Cand. scient. the­ sis. Zoological Museum, University of Bergen. Saaristo, M. I. 1973. Taxonomical analysis ofthe type­ species ofAgyneta, Anomalaria, Meioneta, Aprola­ gus and Syedrula (Araneae, Linyphiidae). Ann. Zool. Fenn. 10, 451 - 466. Schikora, H.-B. 1993. Meioneta mossica sp. n., a new spider close to M. saxatilis (Blackwall) from north­ ern and central Europe (Araneae: Linyphiidae). Bull. Br. Arachnol. Soc. 9, 157 - 163. Scbikora, H.-B. 1995. Intraspecific variation in taxo­ oomic characters, and notes on distribution and habi­ tats ofMeioneta mossica Schikora and M. saxatilis (8lackwall), two closely related spiders from north-

99 ·~,. DII'" ~ t '\ .. .. i' , t ~ "',... t f INSTRUCTIONS TO AUTHORS White 1998). When a paper has more than two authors, it should be referred to as Smith et al. The language is English or occasionally Norwe­ (1990). Multiple references in the text are given gian with an extended English summary. in chronological orders (Smith 1975, Green 1980, Black & White 1998). All references (but not any Manuscripts, double spaced on one side of the that has not been cited in the text) should be listed paper must be submitted in duplicate. Separate in alphabetical order at the end of the paper. In sheets should be used for (1) title page with au­ English reference lists, 0 is equal to 0 and A is thors names, (2) abstract followed by the name(s) equal to Aa. Names of journals are abbreviated and postal address(es) of the author(s), (3) tables, according to international standards, e.g. as in (4) numbered figures and (5) legends to figures. BIOSIS (Biological Abstracts). Acknowledgements should be gathered under a single heading at the end of the text. All manu­ Examples: scripts will be considered by referees before ac­ ceptance. Journal papers Chant, D.A. & McMurtry, lA 1994. A review of Abstract should not exceed 300 words and should the subfamilies Phytoseiinae and Typhlodro­ cover the main results and conclusions of the pa­ rninae (Acari: Phytoseiidae). Int. 1. Acarol. 20, per. A list of up to five key words may be added on 223-310. a separate line below the abstract. Book Tables are typed on separate sheets and numbered Borror, DJ., Tripletorn, C.A. & lohnson, N.F. consecutively. Each table must have a heading. 1989. An introduction to the study of insects. Write "Table" in full both in the text and table Sixth edition. 875 pp. Saunders College Pub­ heading. Avoid vertical lines in the table. lishing, Philadelphia

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Localities. In faunistic papers the names of Nor­ wegian localities should be according to K.A. 0kland (1981), Fauna 34, 167-178, and prefer­ ably the EIS number should be added in brackets.

References. Citations in the text should be writ­ ten as Black (1992), (White 1995) or (Black & CONTENTS Norw. J. Entomol. 47 (1) 2000 ISSN 1501-8415.

Articles

Kobro, S., Teksdal, A. E. & Andersen, A. Cereals as host plants for thrips (Thysanoptera) in Norway I 0degaard, F. & Ligaard, S. Contribution to the knowledge of Norwegian Coleoptera i Fjellberg, A. & Jucevica, E. A new species ofArchisotoma Linnaniemi, 1912 from the Baltic coast of Latvia (Collembola, Isotomidae) 21 Gogstad, G.O. Acid rain and the disappearance of the apollo butterfly (Parnassius apollo (L., 1758» from coastal areas in Norway 25 Andersen, I, Olberg, S. & Haugen, L. Saproxylic beetles (Coleoptera) ofTroms and western Finnrnark, northern Norway with exceptional distribution in Fennoscandia 29 Coulson, SJ. A review of the terrestrial and freshwater invertebrate fauna of the High Arctic archipelago of Svalbard ~ I Haenni, I-P. & Greve, L. New records of Norwegian Scatopsidae (Diptera) 65 Hauge, H. & Hansen, L.O. New records of spiders (Araneae) from the Oslofjord area, SE Norway ;3 Aakra, K. New records of spiders (Araneae) from Norway with notes on epigynal characters of Philodromus fuscomarginatus (De Geer) (Philodromidae) and Araneus sturmi (Hahn) (Araneidae) ..,..,

Greve, L. New records of Norwegian Lauxaniidae (Diptera) 89 Aakra, K. Agyneta mossica (Schikora, 1993) (Araneae, Linyphiidae) in Norway 95

Short communications

Olsen, K.M. Mitostoma chrysomelas (Hermann, 1804) (Opilionida, Nemastomatidae), a harvestman new to Norway 24 Olsen, K.M. Haplophilus subterraneus (Shaw, 1789), a centipede (Chilopoda, Geophilomorpha) new to Norway 63 Greve, L. & Johanson, K.A. Malacomyia sciomyzina (Haliday, 1833) (Diptera, Coelopidae) recorded from Norway 72 Hansen, L.O. The family Signiphoridae (Hymenoptera, Chalcidoidea) in Norway 76

Book reviews • Bokanmeldelser , ,., 19

The present journal is a continuation of Fauna norvegica Series B (Norwegian Journal of Entomol­ ogy). With Volume 45 (1998) the publication of this journal by the Foundation for Nature Research and Cultural Heritage Research (NINA-NIKU) has been discontinued. Starting with Volume 46, the journal is published as Norwegian Journal of Entomology by the Norwegian Entomological Society. Editor is Professor Lauritz Somrne, University of Oslo.