SER. B VOL. 32 Orwegian Journal of Entomology

o. 198

SER. B VOL. 32 orwegian Journal of Entomology

PUBLISHED BY ORSK ZOOLOGISK TIDSSKRIFfSE RAL OSW Fauna norvegica Ser. B Norwegian Journal of Entomology Norsk Entomologisk Forenings tidsskrift

Appears with one volume (two issues) annually 75, -. Disse innbetalinger sendes til NZT, Zoologisk Utkommer med to hefter pr. ar Museum, Sarsgt. 1, Oslo 5. Editor-in-Chief (Ansvarlig redaktor) Postgiro 2 34 83 65. John O. Solem. University of Trondheim, The Mu seum, Erl. Skakkes gt. 47, N-7000 Trondheim. FAUNA NORVEGICA B publishes original new in formation generally relevant to Norwegian entomo Editorial Committee (Redaksjonskomite) logy. The journal emphasizes papers which are main Arne Nilssen, ZooligicaJ Dept., Troms0 Museum, ly faunistical or zoogeographical in scope or con N-9000 Troms0, Ole A. S

Members of Norw.Ent.Soc. will receive the journal J free. Membership fee N.kr. 80,- should be paid to the NORSK ENTOMOWGISK FORENING Treasurer of NEF: Lise Hofsvang, Brattvollveien ser sin oppgave i a fremme det entomologiske stu 107, Oslo 11. Postgiro 5 44 09 20. Questions about dium i Norge, og danne et bindeledd mellom de in membership should be directed to the Secretary of teresserte. Medlemskontingenten er for tiden kr. 90, NEF. Trond Hofsvang, P.O. Box 70, N-1432 As pr. M. Henvendelse om medlemskap i NEF sendes NLH. Members of NOF receive the journal by sekret

Norsk zoologisk tidsskriftsentral (NZT) er et felles Managing Editor (Administrerende redaktor) publiseringsorgan for NEF og NOF i samarbeid med Edvard K. Barth, Zoologisk museum, Sars gt. 1, de zoologiske avdelingene ved universitetsmuseene i 0562 Oslo 5. Oslo, Bergen, Trondheim og Troms0. Adresse: Zoologisk museum, Sars gt. 1, 0562 Oslo 5. Editorial Board (RedaksjonsrAd) Postgiro 2 34 83 65. Wim Vader, Troms0, Svein Haftorn, Trondheim, Rolf Vik, Oslo.

Kristiansen & W0ien, Oslo. ISSN 0332-7698

Fauna (Norsk Zoologisk Forening) har gatt ut av Norsk Zoologisk Tidsskriftsentral. Avtalen om gjensidig reduserte abonnementpriser pA foreningens tidsskrifter vii for fremtiden derfor bare gjelde mellom Norsk Entomologisk Forening og :'Ilorsk Ornitologisk Forening. New records of Norwegian Coleoptera 11: Species new to the fauna and notes on some little known species

TORSTEIN KVAMME

Kvamme, T. 1985. New records of Norwegian Coleoptera II: Species new to the fauna and notes on some little known species. Fauna norv. Ser. B, 32,49-51.

Melanotus erythropus (Gmelin, 1790), Apion sanguineum (Degeer, 1775) and Neosirocalus pyrrhorhynchus (Marsham, 1802) are reported new to the Norwegian fauna. Trogoxylon impressum (Comolli, 1837) is reported intercepted to Norway and registrated to oak-wood in a house. New faunal divisions are presented for Stenolophus mixtus (Herbst, 1784) and Coly dium elongatum (Fabricius, 1787). The occurrence of Hylastes angustatus

Torstein Kvamme, Norwegian Forest Research Institute, p.a. Box 61, N-1432 As-NLH, Norway.

Below are presented species, which not previo usly have been reported from Norway. In addi tion informations on some rare or little known species are discussed. The faunal codes are in ag reement with 0kland (1981), and the EIS-grid numbers are added in parenthesis. The nomenc lature follows Silfverberg (I979).

SPECIES NEWTOTHE FAUNA Elateridae Melanotus erythropus (Gmelin, 1790) (rujipes Herbst, 1784). One male and three female were sampled from an about 4 m. tall Ulmus glabra Huds. at VE:Stavern (EIS: 19), 23. June 1981. M. erythropus is very similar to M. castanipes (Paykull, 1800), and was treated as one species by Lindroth (I960) and others. Lohse (I 976) concluded that they are two different species. Palm (1978) has studied the species in Sweden and found that M. erythropus is rare and probably restric ted to the southern and central parts of Sweden, while M. castanipes is widely distributed and com mon. Fig. I shows the distribution of M. castanipes in Norway. The Norwegian material of Melanotus needs an examination to check whether specimens of M. erythropus and M. castanipes have been mixed.

Lyctidae Trogoxylon impressurn (Comolli, 1837). This south-palaearetic species was caught in great numbers at AK:Frogn, S0ndre Hallangen (EIS:28), 6. June 1983. The beetles swarmed in a summer cot tage, and appeared to have hatched from stems ofoak Fig. I. The distribution of Melanotus castanipes (Pay (Quercus robur L.) used ornamentally. kul1) depicted on EIS-grid map.

Fauna norv. SeT. B, 32,49-51. Oslo 1985. 49 The history of introductIon is "classical»: The ow atum was first reported from AAY:Ris0r (EIS: 11) ner of the cottage spent his vaccation in Northern (Pasche & Zachariassen 1976). In June 1981 more Italy 6-7 years ago. He brought back some nice than 25 specimens were caught in pipe-traps and wood-pieces of grapevine as a souvenir. When speci window-traps at VE:Brunlanes, Solbergstua mens of T. impressum from the grapevine hatched, (EIS: 18). The traps were placed at a sun-exposed they established in oakstems. The cottage is not artifi clearcut. The clearcut was surrounded by stands of cial heated in long periods of the year, but still the be Norway spruce (Picea abies (L.) KarsU and mixed etles survived. forest. The spruce stands were heavily attacked by T. impressum is known to live in wood of grape [ps typographus (Linnaeus, 1758). Two specimens of vine, fruit trees and poplar (Freude et al. 1969). Ear C. elongatum were also sampled from a window-trap lier, T. impressum has been introduced to Denmark at B0:Hurum, Filtvedt (EIS:28) (Leg. F. Midtgaard) (Silfverberg 1979). in the period I. July to I. September 1983. C. elonga tum is living under the bark of both conifer and deci dious trees attacked by bark beetles (Palm 1959, Han Apionidae sen 1964). The heavy attacks by I. typographus has Apion sanguineum (Degeer. 1775) (miniatum Ger probably made temporary optimal conditions for C. mar, 1833). elongalUm. A revision of the subgenus Erythrapion showed that former reports of A. sanguineum from Norway were based on misidentifications (Kvamme 1981). One Cerambycidae specimen of this species was found in a pit-fall trap Agapanthia villosoviridescens (Degeer, 1775). on the little island Skurvene in AAY:His0Y (EIS:6l, 4. This longhorn beetle is usually regarded to be rare, August- 23. September 1981 (Leg. Torstein Solh0Y). and is found only sporadically in South-Eastern Nor A. sanguineum is living in the rootcollar and the lo way. The larva is observed in different plants: Eupa wer parts of the bigger species in the genus Rumex t6rium canndbinum L., Circium olel)llceum (L.), and (Dieckmann 1977). Arclium IOmenlOsum Mill. (Palm 1953). Horion (1974) mention also Anthriscus, Angelica, Carduus, Chaerophyllum, Heracleum, Senecio, Urlica and Curculionidae especially Circium arvense (L.) Scop. and C. palustre Neosirocalus pyrrhorhynchus (Marsham, 1802). (L.) Scop. as hostplants. At HES:Ringsaker, Mengs One specimen was found by sweep-netting Sisymb hot (EIS:45) were eleven specimens sampled from. rium sp. in AK:Oslo (EIS: 28), 4. June 1982. This is an the lower parts of the stalk of Aconitum septentrio expected species, while N. pyrrhorhynchus is known nale Koelle, 23. May 1981 (Leg. L. Aarvik). One spe from several faunal divisions in South-Sweden (Lin cimen was also sampled under the same condition at droth 1960). HES:Stange, Rotlia (EIS:46), 3. June 1981 (Leg. L. Aarvik). The beetles were found as pupae or newly hatched imago, and these finds represent a new NOTES ON LITTLE KNOWN SPECIES faunal division besides a new host. Carabidae Stenolophus mixtus (Herbst, 1784) has expanded Scolytidae northwards in this century, and was first recorded in Hylastes angustatus (Herbst, 1793). Norway in 1977 (Kvamme 1978). Two new localities The opinion on the occurrence of H. angustallls in have been found: Five specimens were sampled at Norway has been divided. Lekander et al. (J 977) VE:Hedrum, Roppestad (EIS: 19), 5. June 1979. One have deleted this species from the Norwegian list in specimen was also recorded at AK: Vestby, Tjenn their revision of the Scandinavian Scolytidae. They (EIS:28), 19. June 1979. Both records were made have included the finds of H. angustatus in H. allenu close to the water-line among dead plants on muddy allls Erichson, 1836. Strand (see Lindroth 1960) soil. mention H. angustatus from 0, AK, TEy, AAy and Ry. Fjellberg (1966) has published a find from YE.

Colydiidae I have seen two specimens of H. angustatus from Colydium elongatum (Fabricius, 1787) and C. flU- VE:Sandar (EIS: 19) (Leg. A. Vik) determinated by forme Fabricius, 1792. . Strand. These specimens agree with H. angustatus The two species belonging to the genus are rare in sensu Grocholski et al. (J 976). Since Strand's identifi Norway. C. filiforme is only known from cations were correct, I find it correct to include the TEY:Porsgrunn, Brevik (EIS: 11). The only specimen finds examined by him. Specimens from 0, AK, VE is stored in Museum of Zoology in Bergen. This re and TEY have been examined by Strand (Strand cord was made in the last century (Ex coil. N.G. pers.com.). Other records made by Munster and Hel Moe), and the species is not found later. The occur liesen must be verified before they are included in the rence in Norway today needs to be verified. C. elong- list.

50 AKNOWLEDGEMENTS Kvamme, T. 1981. The red Apion species in Norway (Col.. Apionidael. Fauna 110rv. Ser. B. 28. I would like to thank Leif Aarvik and Fred 35-38. Midtgaard, As, and Anders Vik, Sandefjord for Lekander, B., Bejer-Petersen, B., Kangas, E. & sending me specimens and data. Bengt Ehn Bakke, A. 1977. The Distribution of Bark Beetles strom, Uppsala, and Jyrki Muona, Oulo, have in the Nordic Countries. Acta el1t..fel1l1. 32. 1-37 been very helpful and controlled the determina + maps. tion of Hylastes angustatus and Melanotus eryt Lindroth, CH. (Red. Cur.l1960. Catalogus Coleopte hropus. rorum Fel1noscandia et Dania. EntomoJogiska Siillskapet i Lund. Lohse, G.A. 1976. Elateriden-Studien. EI1I. Blatter REFERENCES 72. 90-104. Dieckmann, L. 1972. Beitriige zur Insektenfauna der Palm, T. 1953. Anteckningar om svenska skalbaggar. DDR: Coleoptera-CurcuIinidae: Ceutorhynchi 7. Ent. Tidskr. 74, 7-23. nae. Beitr. Ent. 22. 3-128. Palm, T. 1959. Die Holz- und Rindenkiifer der slid Fjellberg, A. 1966. Koleopterologisk bidrag til Vest und mittelschwedischen Laubbiiume. Opusc. Ent. folds fauna n. Norsk ent. Tidsskr. 13.144-154. Supp!. 16. 1-374. . Freude, H., Harde, K. W. & Lohse, G.A. 1969. Tere Palm. T. 1978. Melanotus rufipes Herbst. och M. cas dilia, Heteromera. Lamellicornia. Die Kaler Mit tanipes Payk - taxonomisk status och utbred teleuropas 8. 1- 388. ning (Col., Elateridael. Eot. Tidskr. 99. 49 - 51. Grocholski, J., MichaIski, J. & Nowak, W. 1976. Pasche, A. & Zachariassen, K.E. 1976. Coll'diul11 Notes on Intraspecific Variation and Sexual Di elongatum Fabr. (Col., Colydiidael new to Nor morphism of Some PaIearctic Species in the way. Norw. J. Ent. 23. 207. Genus Hvlastes Er. (Co!.. Scolvtidae). Acta Zool. Silfverberg, H. (Red. Cur.l 1979. Eoumeratio Coleop Cracovie~sia 21. 553-584. . terorum Fennoscandiae et Daniae. Helsingfors Hansen, V. 1964. Fortegnelse over Danmarks Biller Entomologiska BytesfOrening. . (Coleopteral Ent. Meddr 33. 1-507. 0kland, K.A. 1981. Inndeling av Norge til bruk ved Horion, A. 1974. Cerambycidae - Bockkiifer. Fau biogeografiske oppgaver - et reviderl Strand-sy nistik der Mitteleuropaischen Kaler 12. 1-228. stem. Faul1a 34, 167 -178. Kvamme, T. 1978. Stenolophus mixtus Hbst., and ex panding Carabid beetle new to Norway. Norw. 1. Received 18 Sep. 1984 El1t. 25. 227-228.

51 Agonum dorsale (Pontoppidan) (Col., Carabidae), an expanding species in Norway

ARILD ANDERSEN

Andersen, A. 1985. Aganum darsale (Pontoppidan) (Col., Carabidae), an expanding species in Norway. Fauna nary. Ser. B. 32, 52-57.

Agonum dorsale has expanded its distribution in Norway since it was first reported from the country in 1965. New districts reported are 80, TEY, VAY, VAI and AAI. Its present distri bution covers the EIS grid numbers I, 2,4-6, 9-12, 16-21, 26-29 and 36. The species was found in 40 of I 12 locations, mainly cereal fields, visited in 1983 - 1984. It immigrated into three intensively investigated areas in 0stfold and Akershus counties du ring 1978-1984. A. dorsale is at present often a common beetle in preferred habitats in parts of southern Norway, and still immigrates into new areas. Possible immigration routes into Norway are discussed, and further expansion is hypothesized. The change to cereal monoculture in Nor way during the 1960-ies may partly explain the expansion.

Arild Andersen, Norwegian Plant Protection Institute, N-1432 As-NLH, Norway.

INTRODUCTION (EIS 6) 2 July 1979 (Simonsen pers. commJ The carabid beetle Agonum dorsale is widesp In spite of the seemingly restricted distribu read and very common all over Europe and in tion in Norway before 1980, some of the data parts of Asia and Africa. In Sweden and Finland suggested a wider distribution and that the spe it reaches north to about 60 0 N (Turin et al. cies was expanding. This was first suggested by • 1977). It is a polyphagous predator appearing Kvamme (1977). An expansion of A. dorsale mainly in warm, dry and light agricultural fields into a new area was demonstrated in swede fi (Thiele 1977). By feeding on pest species like ap elds at a farm at JeI0Y, south-east Norway du hids and lepidopterous larvae it is considered a ring 1975- 1981: 0 Moss: Ramberg (EIS 19) highly beneficial insect to agriculture (Basedow (Andersen 1982). et al. 1976, Scheller 1984). A. dorsale hibernates The purpose of the present investigation was in aggregates under stones at the edge of fields, to find the current distribution of A. dorsale in which makes it easy to detect at that time of the Norway. The material, together with older data, year. In spring and early summer the beetles will be analysed to see whether an expansion migrate into nearby fields to reproduce. The lar has occurred or not. vae have been found in July-Aug., and in the autumn a new generation of beetles hatch and leave the fields to hibernate (Kreckwitz 1980). MATERIAL AND METHODS The first Norwegian find was reported from Different investigators caught carabids in pitfall Vestfold county (VE Tj0me: Kjcere (EIS 19» in traps in vegetable fields at a farm located at AK 1965 (Fjellberg 1966). For all locations the revi As: Gardsbruket NLH (EIS 28) during sed Strand-system (0kland 1981) and the EIS 1981-1984. During 1983 -1984 handpicking system (European Invertebrate Survey, Norwe Was used at the farms Moss: Ramberg and As: gian grid numbers, 0kland 1976) are used. Until Gardsbruket NLH and at a farm located at AK 1980 the following locations were added: Ski: Enga (EIS 28) in Sept. - Nov. to decide o Moss: Ramberg (EIS 19) 20 July 1975 (An whether A. dorsale was present or not. dersen 1982); Moss: AIby (EIS 19) 10 April To find the total distribution of the species in 1976 (Kvamme 1977). VE Tj0l1ing: Svinevika Norway, selected fields, mainly cereals, were in (EIS 19) 23 March 1975 (Kvamme 1977); Tj01 vestigated by handpicking at short visits in ling: Bisjord (EIS 19) 25 May 1979 (Borgersen Apr. - Mayor Aug. -Oct. 1983 -1984. A total pers. commJ AAY Grimstad: Grimstad (EIS 6) of 112 fields were visited in 24 EIS grids in before 1970 (Strand 1970); Trom0Y: Brekka southern Norway.

52 Fauna norv. Ser. B. 32: 52-57. Oslo 1985. Table I. Trapping data at Jeloy, As and Ski 1975- 1984. % =percent of total carabid catch

Year Trapping method Trapping period No. of No. of A. dorsale trap days carabids No. % Investigator" Jeloy 1975 Pitfall traps 30 May-IO Oct. 10530 1697 2 0.1 1976 Pitfall traps 31 May-4 Oct. 10206 2776 0 1978 Pitfall traps 8 June-14 Sept. 4410 1012 0 1979 Pitfall traps I June- 29 Aug. 2670 540 25 4.6 1980 Pitfall traps 23 May-29 Aug. 2940 1129 308 27.3 1981 Pitfall traps 12 June - 29 Aug. 1386 767 J97 25.7 1984 Handpicking 13 Sept. 673 As 1978-1981 Pitfall traps May-Sept. 21812 11403 0 1981 Pitfall traps 31 July-17 Sept. 1000 494 0 0strem 19&2, Semb 1982 1982 Pitfall traps 7 July - 17 Sept. 756 151 2 1.3 Kjreraas and Svagard 1983 1983 Pitfall traps 27 May-8 Sept. 624 215 4 1.9 Hofsvang 1984 Pitfall traps 4 June-lO Sept. 582 286 4 1.4 Hofsvang 1983 Handpicking 14 Sept. - 31 Octo 122 1984 Handpicking 12 Sept. 9

Ski 1979-1980 Pitfall traps May-Aug. 13194 1107 0 1983 Handpicking 4 Nov. 0 1984 Handpicking II Sept. 200 "Investigated by present author unless otherwise stated.

During 1979-1980, carabids were caught in RESULTS a cabbage field at Ski (Andersen 1982), but A. dorsale was not present (Table I). Searching for Population increases the species under stones in the same field in The catches ofA. dorsale in swede fields at Jel0Y 1983 gave no specimens, but in 1984 more than in 1975-19.81 are shown in Table 1. In this 200 specimens were found (Table I). It has pro period of time the species increased in abun bably immigrated into the field the same year. dance from a very rare carabid to a dominating one. The location was revisited in 1984, and a total of 675 specimens of A. dorsale were found Distribution in Norway under stones all over the farm (Table I). Obvio Table 2 shows the finds from 1980 or later and usly it was still a dominating species at the farm. completes the list of known locations ofthe spe During 1978 -1981, carabids were caught in cies in Norway. Fig. I shows the known distri three swede fields (up to 1500 m apart) at As bution of A. dorsale in Norway before and after (Andersen 1982), but no specimens ofA. dorsale 1980. Table 3 and Fig. I summarize the samp was caught (Table I). In 1981-1984 various in ling results from the 112 visited locations in vestigators used pitfall traps in vegetables in southern Norway. The species was present in another field at the same farm. As shown in 35.7% of the locations, and it was found in 20 Table I, the species was caught in low numbers of the 24 searched EIS grids. It was often abun from 1982. In 1983 -1984 the three fields in dant in preferred habitats, especially in Tele vestigated in 1978 -1981 were visited again, mark and Vestfold counties and in coastal areas and a total of 131 specimens (2 + 48 + 8I) of 0stfold and Aust-Agder counties. It was were found under stones (Table I). Also at As found up to an altitude of 200 m (AAI Bygland: the species has immigrated recently and establis Lauvdal), and just north of 60 0 N (B0 Ringerike: hed itself as a rather common beetle. Rytteraker).

53 Table 2. Locations of A. dorsale added after 1980.

Geographic EIS-grid No of region number Location Date specimens Collector" 0 12 Halden: R0dsvannet 27 Sept. 1983 69 19 Rygge: Ror 16 Apr. 1984 35 20 Halden: Fredriksten 13 Sept. 1981 2 P. Ottesen Halden: Ysthede 16 Apr. 1983 16 Borge: Arum 16 Apr. 1984 4 Ons0y: M0klegard 16 Apr. 1984 1 Racte: Tom 16 Apr. 1984 29 Skiptvedt: Holstad 16 Apr. 1984 11 Skjeberg: Torsnes 16 Apr. 1984 1 Valer: Nordby 16 Apr. 1984 2 21 Aremark: Fange 16 Apr. 1983 I 29 Spydeberg: Heli 16 Apr. 1984 18 AK 28 Asker: Sem 18 June 1981 1 J.H. Simonsen Oslo: Ekebergsletta 27 Apr. 1982 1 Ligaard 1984 B0 19 Kongsberg: Hvittingfoss 22 Sept. 1983 25 27 Kongsberg: Skollenborg 22 Sept. 1983 5 Flesberg: Svene 21 Sept. 1984 2 28 Hurum: Filtvet 14 May 1982 numerous F. Midtgaard Humm: Tofte May 1982 nume- J rous F. Midtgaard Hurum: Kana 30 Aug. 1983 5 Lier: Tuverud 31 Aug. 1983 1 Lier: Lierbyen 26 Sept. 1983 10 36 Ringerike: Rytteniker 23 Sept. 1983 1 YE 19 Stavern: Rugland 2 May 1980 I T. Kvamme T0nsberg: T0nsberg gym. 19 June 1981 2 S. Ligaard Tj0me: Gon Apr. 1982 numerous A. Fjellberg Tj0me: Havna Apr. 1982 numerous A. Fjellberg Borre: Bast0Y 30 May 1982 2 T. Kvamme Borre: R0re 25 Sept. 1982 4 23 Apr. 1983 85 Sem: Akersmyra 8 act. 1983 25 Andebu: Askjemvannet 20 Apr. 1984 5 Larda1: Hem 21 Sept. 1984 15 28 Svelvik: Knem 30. Aug. 1983 4 TEY 11 Bamb1e: Stokkavatnet 7 Apr. 1983 14 Bamble: Vag 7 Apr. 1983 I 18 Skien: Gjerpen 7 Apr. 1983 68 TEI 16 Tokke: Dalen 19 Sept. 1984 7 17 Kviteseid: Sundkilen 25 Aug. 1982 numerous Ligaard 1984 Seljord: SeIjord kirke 19 Sept. 1984 39 26 Seljord: Flatdal 19 Sept. 1984 3 27 Notodden: Tveiten 22 Sept. 1983 3 AAY 6 Trom0y: Revesand 25 Apr. 1980 I J.H. Simonsen Moland: Brekka 7 Apr. 1983 96 Lillesand: Sangereid 7 Apr. 1983 19 10 Tvedestrand: Lunde 31 May 1984 1 11 Ris0r: Indre S0ndeled 8 Oct. 1983 11 AAI 9 Byg1and: LauvdaI 19 Sept. 1984 9 VAY I Farsund: Hananger 10 act. 1983 13 2 MandaI: Berge 8 Oct. 1983 2 Songdalen: Rosseland 8 Oct. 1983 15 5 Kristiansand: Hamre 7 Apr. 1983 22 VAI 4 Kvinesdal: Slimestad 10 act. 1983 I 20 Sept. 1984 5 "Investigated by present author unless otherwise stated Table 3. Survey of A. dorsale in southern Norway 1983-1984

EIS-grid Negative Positive search number search No. of No. of locations specimens I 4 I 13 2 I 2 17 3 8 o 4 5 I 6 5 o I 22 6 3 2 115 7 8 o 9 I I 9 10 I I I II o 3 26 12 o I 69 16 I I 7 17 o I 39 18 o I 68 19 o 6 169 20 o 7 64 21 I I I 26 2 I 3 27 o 3 10 Fig. I. Distribution of A. dorsale in Norway. 28 12 4 20 • = Known distribution before 1980. 29 10 I 18 • = The present distribution (1984). 35 5 o \l = Searched areas where it was absent in 36 6 I 1983-1984. 37 4 o ------Totals 72 40 678

DISCUSSION In Finland and Sweden A. dorsale has been pre expansion in distribution and commoness must sent north to about 600 N for a long time (Lin have taken place during the last 20 years. droth 1945). In Finland no recent expansion has A. dorsale is always macopterous (Lindroth been observ~d (Silfverberg pers. comm.). In 1945) and has been observed flying (Kreckwitz • Sweden the species has become more common 1980). This makes migration by flight a reaso in later years and it was caught new to Dalsland nable hypothesis, which is strengthened by the in south-western Sweden in 1982 (Baranowski sudden appearance of the species in new areas pers. commJ like at Ski in 1984. Flight is expected to be an A. dorsale was reported from Norway for the important factor in the rapid expansion of A. first time in 1965 and today it is widespread in dorsale in Norway. the area shown in Fig. I. It is not easy to recapi The most probable immigration route into tulate the pattern of expansion, but a few re Norway is from south-western Sweden to 0st marks will be done. The species appears in one fold county (Fig. 2). If so, the Oslofjord may of the most intensively investigated areas of have acted for some time as a barrier for further Norway as to entomology (Kvamme 1977), and expansion westwards. The species is uncommon it is also easily recognized due to its typical colo around the northern parts of the fjord, but has urs. Accordingly it is unlikely that the species probably crossed the fjord at narrower parts (Je have been a fairly common beetle here for long 10y (0) - Horten (YE): 5 km, Df0bak (AK) without being discovered. However, since it is a Hurum (B0): 1.5 km). An alternative immigra common species in such a large area today, the tion route is by wind-drift across Skagerak (I 20 expansion has probably been going on for seve km) from northern Denmark to Aust-Agder ral years. The species may have been rare in county (Fig. 2), as the species has been shown to parts of the area for some time, but most of the be transported in this way about the same dis- 55 Fig. 2. The present distribution (I 984) ofA. dorsale in gaard, F. Midtgaard, P. Ottesen, H. Silfverberg , Fennoscandia and Denmark. and J.H. Simonsen, R. Baranowski, T. Kvarnme • = possible immigration routes into Norway. and P. Ottesen also critisized tl1e manuscript. • = possible further expansion in Norway. Part of the investigation was financially suppor ted by the Agricultural Research Council of tance across the Baltic Sea from the Continent to Norway. the shores ofsouth-eastern Sweden (Baranowski and Gardenfors 1974). A. dorsale is common in areas with human activity, and the species may also have followed plant material, soil etc. im REFERENCES ported into Norway from other European coun tries. Similarly, dispersal by human activity may Andersen, A. 1982. Carabidae and Staphylinidae (CoL) in swede and cauliflower fields in south have occurred within Norway. eastern Norway. Fauna norv. Ser. B 29, 49-61. The. species is very common in cereal crops Baranowski, R. & U. Gardenfors 1974. Vinddrift av (Geiler 195611957), and during the 1960-ies jordlopare i sydostra Skane (Col., Carabidae). En large agricultural areas in Norway changed to tomologen 3, 35-52. cereal monoculture. This has favoured A. dor Basedow, T., A. Borg, R. de Clercq, W. Nijveldt & F. sale and may explain at least part ofits increased Scherney 1976. Untersuchungen tiber das Vor distribution and commonness. kommen der Laufkafer (Col.: Carabidae) auf eu It is presumed that Fig. I gives the correct ropilischen Getreidefeldern. Entomophaga 21, present distribution of A. dorsale in Norway. 59-72. Fjellberg, A. 1966. Koleopterologiske bidrag til Vest However, the species is expected to further ex folds fauna n. Norsk Ent. Tidsskr. 13, 144-154. pand its distribution. The most suitable areas Geiler, H. 1956/1957. Zur Okologie und Phanologie will be westward to Rogaland county (RY, EIS der auf Mitteldeutschen Feldern lebenden Carabi 3 and EIS 7) and to the north in Akershus and den. Wiss. Z. Karl-Marx-Univ. Lpz. 6. 35-61. Buskerud counties (AK, B0 and BV, EIS 35 and Kjreraas, p.a. & U. Svaglird 1983. Effekten av samp EIS 37) (Fig. 2). As the species prefer warm and lanting mellom gulrot og 10k pa gulrotflue (Psila dry areas the expansion in western Norway pro rosae Fabricius). Thesis (cand. agric.), Agricultu bably at last will be stopped by the high humi ral University of Norway (unpublished) 145 pp. dity and in eastern Norway by the long winters. Kreckwitz, H. 1980. Untersuchungen zur Fortpflan zungsbiologie und zum jahresperiodischen Ver halten des Carabiden Agonum dorsale Pont. in ACKNOWLEDGEMENTS Temperatur- und Feuchtigkeitgradienten. Zool. lb. Syst. 107, 183-234. I am indepted to the following persons for data Kvamme, T. 1977. On the distribution and habitat and information: R. Baranowski, B. Borgersen, choice of Agonum dorsale Pont. (Col., Carabidae) A. Fjellberg, T. Hofsvang, T. Kvamme, S. Li- in Norway. Norw. l. Ent. 24, 31-32.

56 Ligaard, S. 1984. L0pebillefunn fra forskjellige deler Scheller, H.V. 1984. The role of ground beetles (Ca av Norge; de fleste fra indre Telemark (TEi) (Cole rabidae) as predators on early populations of ce optera). Fauna norv. Ser. B. 31, 61-62. real aphids in spring barley. Z. ang. Ent. 97, Lindroth, c.H. 1945. Die Fennoskandischen Carabi 451-463. dae. Eine Tiergeographische Studie. I. Spezieller Semb, C.E. 1982. Samplanting. Repellerende stoffer Tell. Goteborgs K. Vetensk. -a. VitterhSamh. fra I0k og naturlige nyttedyr i kontroll av gulrot Handl. Ser. B 4 (I), I -709 + I map. flue (Psila rosae Fabricius). Thesis (cand. agric.), 0k1and, J. 1976. Utbredelsen av noon ferskvanns Agricultural University of Norway (unpublished). muslinger i Norge, og litt om European Inverte 99 pp. brate Survey. Fauna 29, 29-40. Strand, A. 1970. Additions and corrections to the 0k1and, K.A. 1981. Inndeling av Norge tll bruk ved Norwegian part ofCatalogus Coleopterorum Fen biogeografiske oppgaver - et revidert Strand-sy noscandia et Dania. Norsk Ent. Tidsskr. 17, stem. Fauna 34, 167-178. 125-145. 0strem, L.A. 1982. Samplanting. Repellerande stoff Thiele, H.-V. 1977. Carabid beetles in their environ fni bete og naturlege nyttedyr i kontroll av stor og ments. Springer Verlag, Berlin, Heidelberg, New lita kaIflue (Hylemyia jlora/is Fallen og Hylemyia York, 369 pp. brassicae Bouche). Thesis (cand. agric.), Agricul Turin, H., J. Haeck & R. Hengeveld 1977. Atlas ofthe tural University of Norway (unpublished). 119 carabid beetles of The Netherlands. North-Hol pp. land Publishing Company, Amsterdam, 228 pp.

Received 2 Jan. 1985

57 1 Studies ofshortwingedness in stoneflies (Plecoptera)

ALBERT LILLEHAMMER

Lillehammer, A. 1985. Studies of shortwingedness in stonefiies (Plecoptera). Fauna norv. Ser. B. 32, 58-61.

Capnia atra Morton, Amphinemura standfussi Ris and Leuctra hippopus Kempny were ana lysed for shortwingedness. L. hippopus and A. standfussi had a significant relationship between wing length and body length. Small-body populations ofA. standfussi were shortwinged and those of L. hippopus longwinged. The most pronounced shortwingedness occurred in marginal or isolated populations and is probably genetic bound and a result of a selection process in the postglacial periqd.

Albert Lillehammer, Zoological Museum, Sarsgt. I, 0562 Oslo 5, Norway.

INTRODUCTION 8 7 Shortwingedness has been mentioned and dis cussed by several authors and more than one ex planation for the phenomenon has been sugges l'... ted. Hynes (I 94 I) stated, that shortwingedness ~ 4 ./ increases with increasing altitude and Brinck ...c (I949) found that the tendency to shortwinged i 2 Amphinemura atandfuaai ness in Amphinemura standfussi Ris was more abundant in the mountain areas than in the low land of Sweden. Lillehammer (I976) noted that 123456189 a tendency to shortwingedness was fairly com Body length (MM) mon in local populations and that shortwinged 8 populations were mainly found at high altitudes. 7 However, shortwinged populations may occur 6 in lowland localities such as reported from lake 5 Vattem in southern Sweden (Brinck 1949). Miil 4 ler (I978) suggested that especially low tempera 3 ture during the egg stage induced shortwinged 2 ness in Capnia atra Morton. Recent studies by Leuctra hippopua Donald and Patriquin (I983) suggest that short wingedness is genetically bound and that short 123456189 wingedness in three Nearctic Capnia species is related to the lakes' geological age. There are in 8 • 7 • • dications that shortwingedness occurs both oc • ••• casionally and regularly in populations. A study 6 ••• of three species, C. atra, Leuctra hippopus 5 • •• Kempny and A. standfussi, occurring in Fenno • 4 scandia and other parts ofEurope was therefore made. Shortwingedness is normally found in the 3 males of a number of species. Often this short 2 Capnia atra wingedness is so prominent that the wings are less than half of normal size (Micropterous). In 1234567.9 species with great variations in the wing length Fig. I. The relationship between wing length and body length of Capnia atra (A), Leuctra hippopus (B) I) Contribution No. 181 from the Zoological and Amphinemura standfussi (C). The regression ana Museum, University of Oslo. lysis, see Table I.

58 Fauna norv. Ser. B. 32: 58-61. Oslo 1985. A

A. s.

c.o.

Fig. 2. (A) normal and (B) short wings of Amphine mura standfussi and Capnia atra.

in One or both sexes, the reduction is less promi nent (Brachypterous). All the three species stu died are brachypterous. Fig. 3. Capnia atra from the «Devil's Punch Bowl», a small lake 700 m a.s.l. in the Mangerton Mountains, MATERIAL. Ireland (O'Connor, 1978). Fore wing 2.3 mm, hind wing 1.8 mm. The materia~ consisted of 400 9 9 and 300 d d of C. atra, 600 9 9 and 400 d d and 150 nymphs of L. hippopus, 540 9 9 and 350 tion of body length in A. standfussi was follo d d At standfussi collected in different parts of wed by a stronger reduction of wing length and Norway during the period 1966-1979. shortwingedness occurred regularly in small body sized populations (Fig. 1). In L. hippopus the reduction of body length was followed by a RESULTS less prominent reduction in wing length, resul Regular and irregular shortwingedness ting in longwingedness in small-body sized po Specimens from several populations were analy pulations. Shortwingedness occurred regularly sed for the relationship between body length (B) in A. standfussi and irregularly in C. atra and L. and wing length (W) in a simple linear regres hippopus populations. sion. Shortwingedness is then a function of two factors, body length and wing length. A. standfussi and L. hippopus had a significant Form ofreduced wings relationship between wing length and body In all the species studied the reduction of the length, while this was not found in C. atra (Tab. wing length was mainly in the apex ofthe wings 1). However, there were clear differences bet (Fig. 2), often followed by irregularities of the ween A. standfussi and L. hippopus. The reduc- wing ribs. In high altitude populations of A.

L_ Table I. Regression analysis of the relationship between wing length and body length, expressed by a simple regression equation y = a + bx in which a and b are constants. N is number of populations used for measure ments.

Species N a b±95% CL r p C. atra 15 4.059 0.326 ± 0.564 0.304 >0.1 L. hippopus 13 1.983 0.704 ± 0.268 0.856 <0.001 A. standfussi 9 -4.102 1.547 ± 0.533 0.945 <0.001

Table 2. Wing length and percent ofwings with irregular veins in females of Leuctra hippopus from five diffe rent localities situated from the coastal area to the high altitudes of inner fjord areas in western Norway.

Locality NM a.s.l. Wing length Irregular m.m. wing veins -x±sd % Sveio, Costal area 26 30 7.32 ± 0.31 0 Sandeid, Midler 32 30 6.70 ± 0.40 5 Sauda, Inner fjord 31 30 6.47 ± 0.30 6 Sauda, Subalpine 20 500 6.27 ± 0.35 23 Sauda, Lowalpine 45 700 5.89 ± 0.34 46

J standfussi this irregularity was followed by a standfussi, such as in that of the Lovidalen po weakened line in the cubitus area, and a reduced pulation. ability for flight. In some C. atra populations L. hippopus is usually long-winged in margi such as in Ireland (Fig. 3) the reduction of wing nal areas of distribution, and the only short length is very prominent and gives a clear evi winged populations are found where the special dence ofreduced flying ability. The combination temperature isolates the local population from • of reduced wing length and irregularity in wing those of the surrounding areas, such as in the ribs was studied in L. hippopus along a transect lake outlets. of five localities from the coast to the alpine There are differences in the distribution bet areas in western Norway. Normal wings occur ween L. hippopus and A. standfussi. L. hippopus red in the lowland while the reduced wings in mainly occur below the tree line and is not re specimens living at high altitude had a high fre corded above the low-alpine vegetation belt. A. quency of irregularity of wing veins (Tab. 2). standfussi is frequently recorded also in the However, irregularities in wing ribs may also middle-alpine areas where the climate is hard occur in the lowland, e.g. in the western margi and the temperature low. Shortwingedness ofA. nal shortwinged population of C. atra at lake standfussi is mainly recorded in alpine and mar Suldalsvatn where about 80% ofthe population ginal populations and the ability ofactive disper had irregularities in wing veins. sal seems interconnected with functional wing length. In alpine areas shortwingedness might be favourable in order to prevent too large loss ofmature adults, in reducing the active dispersal DISCUSSION ability. Shortwinged populations mainly occur Donald & Patriquin (I983) suggested that the at high altitudes also in C. atra (Lillehammer shortwingedness is genetically bound, and some 1976). In low altitudes the shortwinged popula of the data from this study support this assump tions are sometimes recorded in the margin of tion. The irregular shortwingedness found in C. the distribution, such as at lake Suldalsvatn in atra and L. hippopus is probably a result ofa se western Norway, or in isolated populations out lection pressure from the environment. Howe side the continuous distribution of the species, ver, the gradual reduction of wing length found such as at lake Vattern in Sweden. The mentio in A. standfussi (Lillehammer 1976) may also be ned Irish population, which is also a marginal influenced by environmental factors such as one, was recorded at a lake shore about 700 m food quality and temperature differences. Irre a.s.!. (O'Connor 1978). gular shortwingedness can occur also in A. Muller (I978) suggested that especial low

60 temperatures during the egg stage may induce ACKNOWLEDGEMENTS shortwingedness. If so, the same temperature I am grateful to Lars M. 0degiird for correcting has not influenced L. hippopus and C. atra in the the language and to Hans Olsvik, Karsten Sund same manner in the short river between the and Jan H. Simonsen for technical assistance. I lakes Galten and Istern, in southeastern Nor am indebted to Dr. O'Connor, National Mu way. L. hippopus and C. atra occur as adults at seum of Ireland for the loan of Capnia atra spe the same time, but while L. hippopus is short cimens. winged, C. atra is not. At lake Suldalsvatn in western Norway, the same species occur toget her, and they are both shortwinged. In these REFERENCES two outlets the life cycle of L. hippopus differs Brinck, P. 1949. Studies on Swedish stoneflies (Ple from that of the surrounding populations. Beca coptera). Opusc. Ellt. suppl. 11, 1-250. use of the special water temperature of the out Donald, D.B. & Patriquin, D.E. 1983. The wing lets L. hippopus emerges much earlier here than length oflentic Capniidae (Plecoptera) and its rela tionship to elevation and Wisconsin glaciati~n. what it usually does at the surrounding streams. Call. Ellt. 115: 921 - 926. C. atra, however, occurs in the middle of its Hynes, H.B.N. 1941. The taxonomy and ecology of distribution area at lake Galten, and the outlet the nymphs of British Plecoptera with notes on population emerges simultaneously with the the adults and eggs. Trails. R. ell!. soc. Lond. 91. surrounding local populations. At Suldalsvatn 459-557. C. atra occurs outside the normal climatic distri Lillehammer, A. 1976. Norwegian stoneflies. V. butional area and is isolated from other popula Variation in morphological characters compared tions. to differences in ecological factors. Norw. 1. Ell!. It seems obvious that low temperatures alone 23,161-172. Muller, K. 1978. Vingeutveckling hos Capnia atm cannot explain the shortwingedness. It seems Morton i Abisko-omnidet (Plecoptera). Ellt. necessary that some selection process influence Tidskr. 99, 111-113. the tendency to shortwingedness which occurs O'Connor, J.P. 1978. The stonefly Capnia atm Mor in several populations of the three species such ton (Plecoptera, Capniidae) confirmed as an Irish as mentioned by Lillehammer (I976). species. Elltomologist's Gaz. 29. 156- 158.

Received 2 Jan. 1985

61 Distribution and biology of caddisflies (Trichoptera) in Dovrefjell mountains, Central Norway

JOHN O. SOLEM

Solem, J.O. 1985. Distribution and biology ofcaddisflies (Trichoptera) in Dovrefjell mounta ins, Central Norway. Fauna norv. Ser. B, 32, 62-79.

The caddisfly fauna of the middle, low, and sub-alpine zones, ranging from 1630 m to 870 m a.s.l., in the Dovrefjell National Park, was investigated. Only three species, Apatania zo nel/a Zetterstedt, Chaetopteryx vil/osa (Fabricius), and Asynarchus lapponicus (Zetterstedt) inhabited the middle alpine zone, 23 species the low alpine zone, and 60 species the sub-al pine zone. The caddisfly community of the lake outlets in the middle and low alpine zone did not differ markedly from the typical stream fauna, and consisted of limnephilids, except at one lake where the phryganeid Agrypnia obsoleta (Hagen) was present. At the lake outlet in the sub-alpine zone polycentropodids, leptocerids, hydroptilids, and phryganeids, but no hy dropsychids, accompanied the limnephilids. Temporary vernal pools were inhabited by Asy narchus contumax McLachlan, A. lapponicus (Zetterstedt), Grammotaulius signatipennis McLachlan, and Limnephilus stigma Curtis. Pools that have water during the entire warm season, but freeze solid deep down into the bottom substrate in wintertime, were inhabited by A. obsoleta, Oecetis ochracea (Curtis), and Molanna albicans (Zettersteqt). Emergence, Malaise, and light trap data showed that most species peaked early in the flight period. Based on the flight periods the caddisflies can be grouped as follows: I) Late winter species, with adults in April, May; 2) Summer and autumn species, with adults in June to September (R. nubila into October); 3) Autumn species, with adults in late August, September, October. In emergence traps the sex ratio was close to I: I in most species. Signi ficantly more females emerged ofHalesus digitatus (Schrank) and Ecc!isopteryx dalecarlica • Kolenati, and significantly more males emerged of Parachiona picicornis (Pictet) and Plect rocnemia conspersa (Curtis). The dominant feeding group in the middle alpine zone were scrapers, represented by A. zonel/a. Based on dominant species the proportions of sera pers:shredders:predators:collectors in the low alpine zone were 3:3:2:0 and in the sub-alpine birch forest 1:3:3: 1. The parthenogenetic Apatania spp. were dominant in low-competitive early succession habitats which change relatively slowly.

John O. Solem, University of Trondheim, The Museum, Zoological department, Erling Skakkesgt. 47 A, N-7000 Trondheim, Norway.

INTRODUCTION The present paper deals with the caddisfly The first comprehensive, and still the greatest fauna of the surroundings of Kongsvoll in the contribution to the distribution and occurrence Dovrefjell mountains. The area is entirely wit of caddisflies in Norway was given by Brekke hin the Dovrefjell National Park and the adja (I 946). Later, several authors (e.g. Solem 1967, cent protected area. Basic requirements for the I 970a, b, 1977, Tobias & Tobias 1971, Ander erection ofNational Parks in Norway have been sen, T. 1974, 1975, 1979, 1980, 1983, Lilleham topography, vegetation and the composition of mer 1978, Andersen et al. 1978) have given re birds species. One stimulus to the present study ports on caddisflies in selected areas, mostly was the need to increase knowledge ofthe inver from the lowland. But we still have major gaps tebrate fauna in the Dovrefjell National Park. in our knowledge of the occurrence and distri Our National Parks are areas with a high degree bution of caddisflies in Norway, in particular of of protection, and the invertebrate fauna is of vertical distribution. The vertical distribution of great interest in connection with taxonomy, caddisflies at Hardangervidda was reported by long and short term changes of the fauna, and Andersen (I 979), and from the 0vre Heimdalen, aut- and synecology problems. Objectives for Jotunheimen by Lillehammer and Brittain the present study were in taxonomy and aut (I 978). and synecology of caddisflies. Earlier papers on

62 Fauna norv. Ser. B. 32: 62-79. Oslo 1985. Fig. I. Map of the area showing Kallvellsj0el'1 collecting sites.

KOl'1gsvoll ~Iologlcal Statio'"'

Dnva

Isllern

Em"Q"" ,,," 2km ··:· .- Malaise traps . 0 Lrghttrap t Kongsvoll caddisflies which have dealt with selected parts summer season, are inhabited by the same spe ofthe collections for the present paper are Solem cies as the temporary vernal pools, and although (1978, 1979, 1981, 1983a, b, c, d, 1984, 1985). the pools with drought at irregular time periods do not fit exactly into the definition of tempo rary vernal pools, I refer them to this category STUDY AREA because of the caddisfly fauna. The study area was the surroundings of Kongs Two large geological regions in the southern vol1 Biological Station (62°ITN, 09°59'E) bet Scandinavian Calidonides meet in the field area, ween the elevations 870 and 1630 m in the Dov and the border roughly follows the River Driva. refjel1 National Park, Central Norway (Fig. I). On the eastern side is the Trondheim region, The River Driva is the main water course into which contains mainly medium-grade mica which all the smaller streams empty. In general schists and green-stones of cambro-silurian age. the streams are fairly fast-flowing, except for the The western side is mainly a basal gneiss region Jerosbekken. built up of high-grade gneisses and schists of Fol1owing the definitions of the biotic zones precambrian age. in Sj0rs (I967) and R0nning (I972), the samp The climate of the area is mainly continental, ling sites were in three zones. The sampling sites with a yearly precipitation of 473 mm at Kongs of the sub-alpine zone, below 1080 m, were in voU. The yearly mean temperature at Hjerkin the River Driva, the streams Kvernbekken, Rau (959 m a.s.l. and 10 km south of KongsvolI) is bekken, Blesbekken, Vestbekken, Jerosbekken, - 0.1 cC, and only 19 days a year have daily the lake Gavalivatn and temporary pools near mean temperature above 10°C (Nordhagen the JerosbeJqken. In the low alpine zone (1080 to 1943). 1400 m) collections were made at the streams Raubekken, Blesbekken, Kallvel1a and Stropla, the laltes KallveUsj0en and Stroplesj0en, and temporary pools near the Blesbekken. The sam pling sites in the middle alpine zone (above 1400 METHODS m) were the lakes Istjern, Striatjernene, Fiskla The data are mostly based on light, emergence, ustjern and Gluptjern, their outflowing streams, and Malaise trap collections, but also hand net and temporary pools. The highest locality samp col1ections of adults and picking of larvae were led was lake Istjern at 1630 m. periodically made. The numbers oftraps and the There are two types oftemporary pools in the years in which they were used are given in Dovrefjel1 mountains (Solem 1983b)j one conta Table I. Fig. 2 shows emergence and Malaise ins water throughout the entire warm season traps. In the most remote and highest areas, and freezes solid during the winter-time; the se above 1500 m, hand picking of larvae from cond type contains water in spring only, or may rocks and net sampling of adults were the only have short droughts at irregular time periods du methods used. ring the summer season. The type that has water The light, emergence, and Malaise traps were only in spring is termed a temporary vernal pool normally emptied once a week. On a few occa by Wiggins (1973). The pools that have a short sions the most remote traps were emptied at drought at irregular time periods during the two-week intervals.

63 Table I. Number of traps used for adult collections in the Dovrefjell mountains

Light traps Emergence traps Malaise traps Sites 1971 1973 1974 1978 1979 1980 1981 1980 1981 1982 1983 Driva I I 1 Vestbekken 16 Kvernbekken 4 Raubekken 6 8 11 11 2 3 Blesbekken 2 9 5 11 1 3 I lerosbekken 8 Stropla 2 Kallvella 2 Gluptjern 1 Pool A 4 4 Pool B 5 Pool C 3

Fig. 2. Emergence (above) and Malaise traps (below) used.

64 COMMENTS ON TAXONOMIC nella that would justify a distinction into several DIFFICULTIES species, e.g. A. zonella, A. dalecarlica Forsslund or A.forsslundi Tobias. Only 1.47 males per lOO The males and females of Potamophylax cingu females were captured (N =5450) (Solem 1985). latus (Stephens) and P. latipennis (Curtis) are not always easy to assign to species. The females are so much alike that I have made no attempt to DISTRIBUTION OF THE CADDISFLIES distinguish P. cingulatus and P. latipennis when they occurred together. The females have been Middle alpine zone assigned to P. cingulatus and P. latipennis, using The Malaise trap collection at the outlet of lake the proportion of the males as an approxima Gluptjern (Table 2) revealed that A. zonella was tion. A few males appeared with such interme the dominant species. Chaetopteryx villosa (Fa diate characters that they could be assigned to bricius) is also an inhabitant of this biotic zone, either P. cingulatus or P. latipennis, as also re While only one or two individuals of P. cingula ported by Ulfstrand (I970). Thus, there may be tus, Limnephilus coenosus (Curtis), and L. hi a small error in the identifications reported, but punctatus Curtis were collected; they may have this is only a minor obstacle and the main conc been accidental visitors (Table 2). Larvae and lusions of this study are not affected. adults ofA. zonella were collected at lake Istjern Apatania zonella (Zetterstedt) is the only spe at 1630 m, but no C. villosa were found. I can cies captured of the A. zonella-group. Sub-spe not definitely state that C. villosa is not present cies and several species belonging to the A. zo at this altitude, but it cannot be abundant here. nella-group have been described (Svensson & Downstream of lake Istjern, in the area of the Tjeder 1975, Tobias 1981, Malicky 1983), but I lakes Stri

Table 2. Percentage composition of caddisflies collected in Malaise traps in the alpine zone in the vicinity of Kongsvoll biological station

Middle High low Low low alpine zone alpine zone alpine zone Gluptjern Stropla Blesbekken Kall veila Blesbekken Raubekken m a.s.!. 1452 m 1289 m 1350 m 1220 m 1200 m 1100 m Apalania zonella 97.3 16.3 2.1 18.5 1.3 5.7 Chaelopleryx villosa 2.7 5.9 3.0 14.1 0.4 1.6 POlamophylax cingula- IUS , + 14.3 6.2 16.3 3.9 4.4 P. lalipennis 18.8 Limnephilus coenosus + 3.0 3.7 0.5 4.1 Apalania hispida 0.5 11.4 9.8 1.3 2.2 A. muliebris 20.1 0.1 41.9 686 Ecc!isopleryx dalecarlica 33.0 53.7 3.9 24.6 2.5 Rhyacophila nubila 30.0 0.2 14.2 21.4 3.8 Philopolamus mon- lanus 0.2 POlamophylax nigricornis 3.2 5.7 Limnephilus bipuncla- Ius + 0.2 0.8 L. panlodapus 0.1 L. algosus 0.1 Halesus digilalus 0.2 0.1 Apalania sligmalella 0.1 1.0 Beraea pullala 0.2 Microplerna sequax 0.3 Total number of individuals collected 5332 163 568 1016 836 312

65 Table 3. Species recorded in the sub-alpine zone in the vicinity of Kongsvoll biological station

Hydroptilidae Limnephilidae Oxyethira jlavicornis (Pictet) Apatania stigmatella (Zetterstedt) Oxyethira frici (Klapalek) Apatania hispida Forsslund Hydroptila forcipata (Eaton) Apatania zonella (Zetterstedt) Hydroptila tineoides Dalman Apatania muliebris McLachlan Ecc/isopteryx dalecarlica Kolanati Rhyacophilidae Limnephilus algosus (McLachlan) Rhyacophila nubila (Zetterstedt) Limnephilus auricula Curtis Limnephilus bipunctatus Curtis Glossosomatidae Limnephilus borealis (Zetterstedt) Glossosoma intermedia (Klapalek) Limnephilus coenosus (Curtis) Limnephilus extricatus McLachlan Philopotamidae Limnephilus fenestratus (Zetterstedt) Philopotamus montanus (Donovan) Limnephilus femoralis Kirby Limnephilus jlavicornis (Fabricius) Polycentropodidae Limnephilus fuscicornis (Rambur) Plectrocnemia conspersa (Curtis) Limnephilus nigriceps (Zetterstedt) Polycentropus jlavomaculatus (Pictet) Limnephilus pantodapus McLachlan Holocentropus picicornis (Stephens) Limnephilus rhombicus (Linneaeus) Limnephilus sparsus Curtis Psychomyidae Limnephilus stigma Curtis Psychomia pusilla (Fabricius) Limnephilus subcentralis Brauer Colpotaulius incisus (Curtis) J Molannidae Grammotaulius signatipennis McLachlan Molanna albicans (Zetterstedt) Phaecopteryx brevipennis (Curtis) Molannodes tinctus (Zetterstedt) Anabolia concentrica (Zetterstedt) Asynarchus contumax McLachlan Leptoceridae Asynarchus lapponicus (Zetterstedt) Cerac/ea nigronervosa (Retzius) Potamophylax cingulatus (Stephens) Mysticaides azureus (Linneanus) Potamophylax latipennis (Curtis) Oecetis ochracea (Curtis) Potamophylax nigricornis (Pictet) Micropterna sequax McLachlan Beraeidae Halesus digitatus (Schrank) Berea pullata (Curtis) Halesus radiatus (Curtis) Halesus tesselatus (Rambur) Lepidostomatidae Parachiona picicornis (Pictet) Lepidostoma hirtum (Fabricius) Chaetopteryx villosa (Fabricius) Annitella obscurata (McLachlan) Sericostomatidae Chilostigma sieboldi McLachlan Sericostoma personatum (Spence) Notidobia ciliaris (Linnaeus) Phryganeidae Phryganea bipunctata Retzius Agrypnia obsoleta (Hagen)

The temporary pools in the middle alpine (Donovan) were captured in Malaise traps at zone were populated by only one species, A. lap running water. At standing water Agrypnia ob ponieus (Zetterstedt). In total 6 species were soleta (Hagen) was found. found in the middle alpine zone. When defining dominant species as making up more than 20 % ofthe total number collected at one site, E. dalecarlica and R. nubila domina Low alpine zone ted the caddisfly fauna at the Stropla and E. da Twelve species were recored in the upper part of lecarlica and A. muliebris at the Blesbekken the low alpine zone. In addition to the species re (Table 2). corded in the middle alpine zone, Apatania muli In the lower part of the low alpine zone 23 ebris McLachlan, A. hispida Forsslund, Eec/i species were captured. Species additional to sopteryx dalecarlica Kolenati, Rhyacophila nu Table 2 and those mentioned under the high low bila Zetterstedt, and Philopotamus montanus alpine zone, are Grammotaulius signatipennis

66 McLachlan, Limnephilus stigma Curtis, L. fene which are the habitat of B. pullata, but the few stratus (Zetterstedt), and Chilostigma sieboldi specimens ofA. stigmatella have probably flown McLachlan. in from habitats at lower elevation. In the Raubekken A. muliebris outnumbered At Kallvella, a stream on the western side of all other species. A. muliebris, E. dalecarlica, the valley, no particular species dominated the and R. nubila were dominant species from the caddisfly community, but five species, A. zo Blesbekken which runs parallel to the Raubek nella, C. villosa, P. cingulatus, P. latipennis and ken, but only 300 to 400 m away. The Raubek R. nubila each represented between 14 and 19 % ken and Blesbekken are on the eastern side of ofthe total catch. Itshould be noticed that in this the valley. When collecting larvae in the streams stream P. latipennis takes the place of P. nigri ofthe low low alpine zone, Apatania stigmatella cornis (Pictet) in acompanying P. cingulatus. (Zetterstedt) and Beraea pullata (Curtis) were One obvious difference between the Raubekken never found. I did not search in small springs, and Blesbekken, which are inhabited by P. cing-

Table 4. Percentage composition of Malaise trap collections and light trap collections from rapid mountain streams and rivers in the sub-alpine zone in the vicinity of Kongsvoll biological station

Blesbekken Raubekken Jerosbekken Driva river 1000 m 900 m 900 m 870 m Apatania zonella 2.1 1.8 0.3 A. hispida 3.2 A. muliebris 4.4 5.4 Ecclisopteryx dalecarlica 8.8 2.4 0.2 4.2 Potamophylax cingulatus 3.7 21.0 34.6 4.2 P. /atipennis 0.5 11.2 Philopotamus montanus 5.5 1.2 Rhyacophi/a nubi/a 65.2 8.4 20.0 61.5 Chaetopteryx vi//osa 0.2 3.6 3.0 0.3 Limnephi/us coenosus 3.1 20.4 0.2 0.1 Potamophy/ax nigricornis 2.8 7.8 0.1 Limnephilus bipunctatus 0.2 Ha/esus digitatus 0.5 0.4 5.7 Beraea pullata 0.2 Limnephi/us sericeus 0.2 0.1 L. incisus 0.6 P/eetrocnemia conspersa 0.2 0.2 0.1 Apatania stigmatella 1.2 11.2 Micropterna sfquax 4.8 1.7 Parachiona plcicornis 20.4 Limnephi/us sparsus 0.1 L. dispar 0.1 1.2 Ha/esus'radiatus 29.1 0.6 Annitella obscurata 0.1 Po/ycentropus jlavomacu/atus 2.9 0.1 Ha/esus tesse/atus 0.1 0.1 G/ossosoma intermedium 0.1 Hydropti/a tineoides 3.2 Lepidostoma hirtum 2.1 Hydropti/a forcipata 0.7 Oxyethira frici 0.5 Sericostoma personatum 0.9 Limnephi/us fenestratus 0.3 Ceraclea nigronervosa 0.2 H%centropus picicornis 0.1 Anabolia concentrica 0.1 Limnephi/us a/gosus 0.1 L. extricatus 0.1 Total number of individuals collected 619 167 1649 6126

67 ulatus and P. mgrlcornis, and the Kallvella, Table 5. Percentage composition of emergence trap where P. cingulatus and P. latipennis coexist, is collections from rapid streams in the sub-alpine zone that Kallvella is a larger stream. Two new spe in the vicinity of Kongsvoll biological station cies appeared at the Kallvella, Limnephilus pan todapus McLachlan and L. algosus (McLach Vestbekken Kvern lan). bekken Rhyacophi/a nubi/a 33.6 13.2 Plectrocnemia conspersa 21.8 Sub-alpine zone Philopotamus montanus 12.7 2.6 Potamophylax cingulatus 10.9 13.2 Sixty species ofcaddisflies were recorded in this Chaetopteryx vi//osa 10.0 5.3 biotic zone (Table 3). The percentage composi Halesus digitatus 3.6 tion in Malaise traps from different streams is gi Potamophylax nigricornis 2.7 47.4 ven in Table 4 and in emergence traps in Table Apatania mu/iebris 1.8 5. Malaise traps were not used at the Vestbek Other ken and Kvernbekken. Limnephilidae Q Q 1.8 18.4 Only three species R. nubila, P. cingulatus Micropterna and C. villosa were collected from all six streams sequax 0.9 or rivers. The few species lacking in one or two Total number E. dalecarlica, P. montanus, of individuals localities included collected 110 38 L. coenosus, P. nigricornis, Halesus digitatus (Schrank) and Plectrocnemia conspersa (Curtis). All the above mentioned species must be regar ded as widespread in the sub-alpine zone, but cies in trap no I and H. radiatus in trap no 2. In some seem to have a relatively low abundance. trap no I at standing water each of six species The remaining species of Tables 4 and 5 have a represented between 10 and 19 % of the total more restricted distribution, but they may be lo number collected. Additionally, Phryganea bi cally abundant, e.g. Parachiona picicornis (Pic punctata Retzius and A. obsoleta were captured tet) and Halesus radiatus (Curtis). The widesp regularly in net collections close to the lake. read species R. nubila and P. cingulatus are also those that, in general, were most abundant in the streams and the river. C. villosa should pro Temporary pools bably be counted together with R. nubila and P. The warm season pools, which are intermediate cingulatus, because I assume C. villosa to be un between permanent standing water and the tem derestimated in Malaise trap collections. porary vernal pools, contain A. obsoleta, Oecetis ochracea (Curtis), and Molanna albicans (Zetter stedt), while the temporary vernal pools are in Outlets of lakes habited by A. lapponicus, A. contumax McLach The species composition of streams and lake lan, G. signatipennis and L. stigma. Table 7 outlets in the alpine zone did not show great dif shows the percentage composition of the cad ferences. The only difference was that A. obso disfly community in the temporary vernal pools leta, a standing water or slowly-running water located in the low alpine zone and the sub-alpine species, occurred at the outlet of lake Stropl zone. A. lapponicus is the dominating species in sj0en. No leptocerids or polycentropodids occur the alpine zone, while A. contumax is the domi red in the alpine zone. At the outlet of lake Ga nant species in the sub-alpine zone. In tempo valivatn in the sub-alpine zone (Table 6), a num rary pools at 1500 m a.s.l. in the middle alpine ber of limnephilids, e.g. Limnephilus borealis zone only larvae ofA. lapponicus were collected. (Zetterstedt), L. pantodapus and L. rhombicus (Linneanus), belonging to standing water, ap peared together with hydroptilids, e.g. Hydrop A COMPARISON BETWEEN tila tineoides Dalman and Oxyethira!rici (Klapa EMERGENCE AND FLIGHT PERIODS lek), the leptocerid Ceraclea nigronervosa (Retzi Six species occurred in sufficient numbers to al us), the polycentropodids Holocentropus picicor low a comparison and the general trends are nis (Stephens) and Polycentropusjlavomaculatus shown in Figs. 3,4,5 and 6. R. nubila, E. dale (Pictet) and the molannid Molannodes tinctus carlica, and P. nigricornis did not show any dif (Zetterstedt). P. latipennis was the dominant spe- ferences between the emergence and flight peri-

68 Table 6. Number of individuals of caddisflies captured in Malaise traps at the outlet of lake GftvaJivatn and at the outflowing Jerosbekken. M = males, F = females

Slow flowing water Rapid flowing water at outlet downstream the outlet Trap I Trap 2 Trap 3 Trap 4 M F M F M F MF Hydropti/a tineoides 13 2 9 42 H. forcipata 2 8 1 Oxyethira frici 2 2 2 2 2 Limnephi/us rhombicus 25 39 7 13 I 2 L. extricatus 2 23 25 71 I L. coenosus 6 2 9 3 I 2 2 L. borea/is 13 58 3 30 3 L. subcentra/is 20 16 8 15 L. fenestratus I 2 4 2 3 L. algosus I L. pantodapus 21 7 17 6 Potamophylax cingulatus 3 29 34 189 198 98 82 P. latipennis 43 65 14 19 2 6 Halesus radiatus 40 18 177 49 179 77 164 58 H. digitatus I 3 4 I H. tesselatus I I Chaetopteryx vi//osa 30 34 25 18 26 14 8 2 Anabo/ia concentrica I Apatania zonel/a Ecc/isopteryx dalecar/ica I 2 Cerac/ea nigronervosa 18 3 49 33 I Holocentropus picicornis I 1 I I Polycentropus flavomaculatus 26 42 2 14 2 15 3 31 Plectrocnemia conspersa I 2 I Rhyacophi/a nub/la 14 6 31 33 87 42 128 71 Mol/anodes tinctus 4 • Sericostoma personatum 9 5 Lepidostoma hirtum 3 7 23

Table 7. Percentage relative abundance ofcaddisflies in Malaise traps at temporary pools in the alpine and sub alpine zones in the Dovrefjell mountains

Sub- alpine Alpine zone zone Pool A Pool A Pool B Pool C 1979 1980 1980 1980 Asynarchus lapponicus 82.4 84.9 78.7 13.9 A. contumax 66.7 Grammotau/ius signatipennis 16.0 9.7 3.3 6.3 Limnephi/us coenosus 2.5 4.3 16.7 L. stigma 13.2 L. fenestratus 0.7 Agrypnia obsoleta 1.0 0.7 Total number of individuals collected 119 93 150 288

69 Rhyacophila nubila Fig. 3 ~ Ecclisopleryx dalecarlica Fig. 4 Q) .0 E Emergence Iraps 1978 to 81 ::LEmo"o",o "eO' 1078 " 51 ::l ! Z 10

5 1219262 9 July Aug. 1926 2 9 162330 5 132027 80 July Aug. Sept.

600 60 Malaise traps Malaise traps 1980 and 81 500 1980 10 83 '"Q; 40 .0 E ::l 400 Z 20

300 51219262 9162330512 July Aug. Sept. '"Q; 200 .0 Fig. 4. Number of males (white) and females (black) E ::l of Ecc/isopteryx da/ecar/ica captured in emergence Z 100 and Malaise traps. m I 121926 2 9 1623305• 1320274 1118 150 July Aug. Sept. Oct. Apatania muliebris Fig.6 Emergence traps 1978 to 81 Fig. 3. Numbers of males (white) and females (black) '"Q; 100 .0 of Rhyacophi/a nubi/a captured in emergence and E ::l Malaise traps. Z 50

Potamophylax nigricornis Fig. 5 212851219262 91623305 June JUly Aug. Sept. '"Q; 20 .0 E Emergence traps 250 ::l Z 10 19781081 Malaise Iraps 1980 to 83

512192629 July Aug. 150

Malaise traps '"Q; .0 1980 to 83 E ::l iz 2°l10 z50

-5-'--1'21'""-1-1'9"2""6--..J2--9·"':1~6~2'-3.L..3""0 2128 5 121926 2 9 162330-''''''5 July Aug. June July Aug. Sept. Fig. 5. Number of males (white) and females (black) Fig. 6. Numbers of females of Apatania mu/iebris of Potamophy/ax nigricornis captured in emergence captured in emergence and Malaise traps. and Malaise traps.

ods, and the peak ofindividuals occurred simul peak in numbers appeared two weeks later in taneously in the Malaise and emergence traps. the Malaise traps. A. muliebris is a parthenogen One species, A. muliebris, had a different pat tetic species, so there is no need for it to fly aro tern (Fig. 6). The emergence and flight periods und searching for a mate. A female in captivity, covered approximately the same time, but the kept at a temperature of 10°C, oviposited bet- 70 Table 8. Percentage composition of the various stage methods. C. villosa was the only species that in the maturation of eggs in Apatania muliebris seems likely to be underestimated in the Malaise traps. This may be because C. villosa adults do Emergence Malaise not fly, but rather walk on the ground, and the traps traps refore its trapability in Malaise traps is less than for flying insects. This particular behaviour will Small eggs 49.4 5.8 Medium sized eggs 9.2 8.7 not influence the catches in the emergence traps, Large eggs 40.2 52.4 which will therefore give a better estimate of the Spent or partly proportion of C. villosa. This have been taken spent females o 33.0 into consideration when dominant species are Total number of listed in relation to trophic ecology. The num females 87 103 bers of species collected were 12 in the emer gence traps and II in the Malaise traps. ween 2 and 3 weeks after emergence. This diffe rence in time between emergence and oviposi tion coincides with the difference between the FLIGHT PERIODS peak in numbers in emergence and Malaise The flight periods ofthe species captured in Ma traps. It is likely that specimens of A. muliebris laise traps at permanent waters are shown in rest quietly for about two weeks and that the ac Fig. 7; and the total flight period extended from tivity recorded by the Malaise trap represents early June to mid October. The first species to the oviposition flight (Table 8). appear was B. pullata, followed by Apatania hi spida, A. zonella and A. muliebris. Five species were recorded as late as October, R. nubi/a. H. A COMPARISON BETWEEN RELATIVE digitatus, H. radiatus, C. villosa and Limnephi ABUNDANCE AND NUMBER OF Ius subcentralis Brauer. The shortest flight peri SPECIES IN EMERGENCE AND ods lasted only two weeks, as shown by Sericos MALAISE TRAPS toma personatum Spence, Hydroptila jorcipata The percentage composition ofthe caddisflies in (Eaton), H. tineoides, Oxyethira jrici (Klapalek), emergence and Malaise trap catches at A. obsoleta and L.jenestratus. The longest flight II - 1200 m a.s.l. from the Raubekken had i periods extended for three months and were ex high degree of similarity (Table 9). A. muliebris hibited by R. nubila and A. zonella. A. muliebris which dominated the fauna, was collected in and L. coenosus had flight periods only a week closely similar percentages by the two collecting or two less than three months. A fairly sharp peak in numbers of captured specimens during the first half of the flight periods was the most Table 9. Speotes and percentage relative abundance common pattern; examples were: E. dalecarlica, in emergence and Malaise traps in a given area of the P. nigricornis, A. muliebris, and H. radiatus Raubekken in the years 1980 - 81 (Figs. 4, 5, 6, 8). A different pattern, where indi viduals appeared over a longer time with no Emergence Malaise sharp peak, was shown by R. nubila (Fig. 3). Of traps traps 37 species recorded in Malaise and light traps, at Philopotamus montanus 2.7 o permanent running and standing water, 8 spe Apatania muliebris 69.5 68.6 cies was recorded in June, 27 in July, 25 in Au A. zonella 1.1 5.7 gust, 16 in September and 5 in October. A. hispida 0.2 2.2 The flight periods ofthe species inhabiting the A. stigmatella o 1.0 temporary vernal pools showed that A. contu Potamophylax nigricornis 3.9 5.7 max and A. lapponicus emerged earliest, follo P. cingulatus 1.4 4.4 G. signatipennis Chaetopteryx villosa 12.1 1.6 wed by and the last to emerge Limnephilus coenosus 0.5 4.1 was L. stigma (Fig. 9). Ecclisopteryx dalecarlica 0.7 2.5 The family Limnephilidae makes up a high Mieropterna sequax 0.2 0.3 proportion of the caddisfly fauna in high nort Halesus digitatus 0.9 o hern latitudes. Of all the collected caddisflies in Rhyacophila nubila 7.1 3.8 Malaise and light traps the Limnephilidae mad~ Total number of up 68.6 % in the area of Kongsvoll. The tempo individuals collected 439 312 ral distribution of Limnephilidae and non-Lim-

71 June July Aug. Sept. act. 2 7 1421 285 12 1926 29 1623305 1320274 11 15 Apatania muliebris N-737 A. zonella N-5450 A. hispida N-174 A. stigmatella N-626 Ecclisopteryx dalecarlica N-517 Potamophylax latipennis N-564 P. nigricornis N-109 P. cingulatus N-826 Halesus tesselatus N-2 • • H. radiatus N-839 H. digitatus N-309 Micropterna sequax N-13 Parachiona picicornis N-42 Chaetopteryx villosa N-389 Limnephilus pantodapus N-52 L. extricatus N-123 L. rhombicus N-84 L. coenosus N-73 L. borealis N-171 L. subcentralis N-59 L. fenestratus N-6 L. algosus N-5 • • L. bipunctatus N-8 L. sericeus N-1 • Philopotamus montanus N-38 Rhyacophila nubila N-4837 Polycentropus flavomaculatus N-134 Holocentropus picicornis N-4 , Plectrocnemia conspersa N-15 - Agrypnia obsoleta N-17 ... Ceraclea nigronervosa N-107 ~ Sericostoma personatum N-14 Lepidostoma hirtum N-36 - Beraea pullata N-4 • Hydroptila forcipata N-11 ~ H. tineoides N-66 .... Oxyethira frici N-lO .... Fig. 7. Flight periods of species collected in emer gence traps I978 to 1981 and Malaise traps I980 to 1983 from permanent water. Vertical bars denote 50 % of total catch. the second half of August and first half of Sep tember, did non-limnephilids dominate the fauna. The non-limnephilids were heavily domi nephilidae is shown in Fig. 10. The limnephilids nated by R. nubi/a. represented more than 50 % of the flying cad The temporal distribution at Dovrefjell was disflies in June, July, early August, late Septem different from that reported from South Sweden ber and October. Only for about one month, in (Svensson 1972).

72 Fig. 8. Numbers of males (white) and females (black) of Halesus radiatus captured in Malaise traps. 160

120 Halesus radiatus

~ 80 Gl .rJ E £ 40

2 9 162330 5 132027 4 11 Aug. Sept. Oct.

June July Aug. Sept. 1926310172431714212841118 Asynarchus contumax N-192 , A. lapponicus N-236 Grammotaulius signatipennis N-18 , Limnephilus stigma N-38

Fig. 9. Flight periods of species living in temporary vernal pools.

80 SEX RATIOS During the years 1978 to 1981 15 species were 6 identified from emergence traps, and 14 ofthem also from Malaise traps (Table 10). The only spe cies captured only in an emergence trap, was the Limnephilidae single male of Glossosoma intermedium (Klapa 40 lek). No'males of Apatania species (A. hispida. A. muliebris. A. zonella) were collected in emer gence or Malaise traps at these sites. P. mon 20 tanus, P. nigricornis, P. cingulatus, L. coenosus, M. sequax McLachlan, C. villosa, and R. nubila did not show any significant differences bet ween the numbers of males and females captu red in emergence traps, but, except for L. coeno 212851219262 91623305132027415 sus. the numbers ofemerging males were a little June July Aug. Sep!. oc!. higher. Significantly more females than males Fig. 10. Temporal percentage composition of Lim emerged of H. digitatus and E. dalecarlica. nephiIidae species and non-Limnephilidae species ba while significantly more males than females sed on numbers. emerged of P. picicornis and P. conspersa.

73 Table 10. Numbers of male and female caddistlies recorded in emergence traps during 1978 to 1981 and Mala ise traps in Raubekken and B1esbekken 1980 to 1981. * =p <0.5, ** =p <0.01, *** =p <0.00 I, X2-test

Emergence traps Malaise traps Males Females Males Females Philopotamus montanus 26 18 19 21 Apatania hispida 7 33 A. zonella 8 47 A. muliebris 354 712 Potamophylax nigricornis 48 44 77** 48 P. cingulatus 49 44 80 83 Halesus digitatus 3 10 2 5 Parachiona picicornis 13* 4 9 33*** Limnephilus coenosus I 3 45 55 Ecclisopteryx dalecarlica 14 27* 136 160 Micropterna sequax 5 4 7 4 " Chaetopteryx villosa 82 74 7 6 Rhyacophila nubila 90 73 374 397 Glossosoma intermedium I 0 0 0 Plectrocnemia conspersa 25** 7 11 * 2

J TROPHIC ECOLOGY Dominant species have a powerful effect on the ders and gatherers ofsuspended or loosely depo community (Krebs 1972), and when trophic sited fine organic particles; shredders fed on de ecology is concerned, it is therefore legitimate to composing vascular plants; predators were car look at the feeding habits of the dominant spe nivores whose main food items was animal cies only. The abundance of each species varies prey. It seems that most of caddisfly larvae can from stream to stream (Tables 2,3,4,5), but the be grouped under one or another of these hea general trend in the dominance hierarchy is dings. clear, and the dominant species have been grou The Apatania species are scrapers (grazers) ped according to their feeding habits in Table 11. (Wiggins and Mackay 1978), their food being Wiggins and Mackay (978) described the largely diatoms and other algae (Nielsen 1942, functional feeding groups ofaquatic insects pro Lepneva 197 I). C. villosa is classified as a shred posed by Cummins (973) as follows: grazers der, because its overall dominant food is plant (scrapers) were animals which nibbled and scra material (Andersen, J. 1984). E. dalecarlica is, ped periphyton and fine organic materials on according to my own analysis of the gut con rocks and vegetation; collectors were fIlter fee- tents of 10 4th and 5th instar larvae, a typical

Table 11. Dominant running water species in the different biotic zones grouped according to their feeding ha bits

Scrapers Shredders Predators Collectors Middle alpine zone ... A. zonella C. villosa Low alpine zone A. zonella C. villosa R. nubila A. muliebris E. dalecarlica P. cingulatus A. hispida P. cingulatus P. nigricornis P. nigricornis Subalpine zone A. stigmatella C. villosa R. nubila P. montanus H. radiatus P. conspersa P. cingulatus P. cingulatus P. nigricornis P. nigricornis

74 Table 12. The proportions of functional feeding categories of caddisflies. The genus is the basic unit for the data from North America, and the dominating species for the data from Norway. North American data taken from Wiggins and Mackay (J 978)

Scrapers Shredders Predators Collectors Western North American mountain forest biome 4.5 4 1.5 Eastern North American mountain forest biome 3 4 2 Norwegian mountain birch forest biome 3 3

shredder. P. cingulatus and P. mgncornis are, DISCUSSION however, difficult to assign to shredders or pre Clear differences in the numbers of species cap dators, because the two species are opportunistic tured and inhabiting the various vegetation in their feeding in the 4th and 5th instars (Solem zones in the Dovrefjell montains were found. 1983a), and the proportion ofanimals in the gut Two species, A. zone//a and C. vi//osa, were content is likely to be underestimated. How dif common in the middle alpine zone; A. lapponi ficult these problems are was demonstrated by cus inhabited such peculiar habitats as tempo Martin and Mackay (I 982) who fed Rhyacop rary pools and thus had more restricted and fe hi/a larvae with known numbers ofeach of two wer habitats than A. zone//a and C. vi//osa, prey taxa (Ephemeroptera: Ephemeridae and which live as larvae in permanent lotic and len Diptera: Chironomidae) in an artificial stream tic waters. Only limnephilids occurred in the under controlled conditions in the laboratory. middle alpine belt, and one of the species, A. zo They discovered that large numbers of heavily ne//a, was also the only caddisfly recorded as far sclerotized fragments of legs, cerci, wing pads, north as 80° on the island ofSvalbard (Solem et head capsules and abdominal segments which al. 1977). are used in the identification of the prey, were Some characteristics of the species occurring cleaned of soft tissue, but not eaten. Soft tissues in the middle alpine zone are: A. zone//a has ma will appear as unidentified matter in the gut and inly parthenogenetic reproduction, its feeding are mostly referred to as detritus. It seems ob habit is scarping «Aufwuchs» on rocks, it flies in vious that the predatory habit of P. cingulatus early summer, the life cycle may extend over and P. nigricornis will be underestimated, and two or more years (Solem 1985), and it inhabits the species are likely to be more dependent on permanent lentic and lotic water; C. vi//osa rep animal food' than can be measured by the food roduce by mating of males and females, it is a analysis method used by Solem (I 983a). I there shredder, adults appear in late summer and au fore record them as both shredders and preda tumn, the life cycle extends over two years, in tors. R. nubila and P. conspersa are typical pre habits lentic and lotic permanent water, and the dators (Siltala 1907, Wallace and Merritt 1980, individuals have not been seen flying; A. lappo Wiggins and Mackay 1978), while the only col nicus, males and females participate in repro lector is P. montanus. Philopotamid larvae feed duction, feeds as a shredder, flies in summer, the primarily on fine detritus and diatoms (Siltala life cycle is one year with very fast larval growth 1907, Wallace and Merritt 1980). in early summer, and inhabits temporary pools. Table 11 shows the dominant species and Judging from the known biological features (So their assignment to the various functional fee lem 1983a, 1985) there is no niche overlap bet ding groups. When compared with Table 12 it ween these species. The reproductive strategy can be concluded that the scrapers predominate and the feeding habit of A. zone//a is probably a in the middle alpine zone. The proportion bet great advantage for a caddisfly to be successful ween scarpers:shredders:predators:collectors in inhabitant of high altitudes and high latitude the low alpine zone is 3:3:2:0, and that in the areas, e.g. the highest waters in the Dovrefjell sub-alpine birch forest is 1:3:3: l. There are re area and on Svalbard. These are low-competi cognizable differences in the proportions of the tive, early succession habitats which change re various functional units between all three biotic latively slowly. These features are found also in zones considered. the parthenogenetic A. mu/iebris and A. hispida,

75 even though the competition with other caddisf In total 60 species were recorded in the Dov lies is greater at these habitats, e.g. Raubekken refjell mountains and this is about one third of and Blesbekken. However, the distribution of the number of species of caddisflies reported the parthenogenetic Apatania spp. agrees with from Norway. From the River Vindelelven in the pattern ofdistribution reported for partheno Swedish Lapland Ulfstrand (I 970) identified 82 genetic weevils (Vepsa!iiinen & Jarvinen 1979). species, and Andersen, T. (I 979) listed 32 spe At lower altitudes the competition between spe cies from the mountain plateau Hardanger cies is greater and populations consist of males vidda. Gothberg (I 970) reported 57 species from and females. the stream Kaltisjokk (North Sweden) and Ma C. villosa is active at night (Andersen, J. 1984) licky (I 978) 45 species from the birch forest in and because it is an autumn species it will be fre the area around Abisko (Swedish Lapland). In quently exposed to temperatures below O°e. In Norway, just west of Vindelelven, 32 species of the year 1982 there was a snowfall in August caddisflies were recorded in light traps (Solem above about 1500 m, and the snow stayed unpubl. data). These data indicate that fewer throughout the autumn. How do individuals caddisfly species belong to the sub-alpine or al cope with these conditions? Do they freeze or do pine zones in South and Central Norway than in they supercool? Supercooling points ofC. villosa Sweden. ranged from -4.5° to -14.5°C (mean It should be noted that no Hydropsyche spe -8.5°C, N= 10)(Solem 1984). This range was cies were collected in this part in the Dovrefjell much wider than that found for summer species mountains, and only one Rhyacophila species of caddisflies, R. nubila, L. borealis and L. sub was recorded. Most of the streams drain glacier centralis, which was -4.0° to -8.9°C (N to areas and low temperatures and )ittle organic tal = 19). C. villosa disappeared in 1982 at the drift may be a reason for the absence of the lar time when minimum night temperatures were vae of Hydropsyche which are filter-feeders. down regularly to - 8°e. Low air temperature In more southern latitudes, e.g. Britain is a limiting factor for adults of C. villosa to stay (Crichton 1960, 1970, South Sweden (Svensson alive in autumn. The mean supercooling point 1972) and Czechoslovakia (Novak and Sehnal of C. villosa is within the range that is recorded 1963, Malicky 1981) some caddisfly species • for species with no special adaption for low tem have two peaks in flight activity during a sea peratures (S0mme & Zachariassen 1981). Howe son. No such feature can be deduced from the ver, the supercooling points indicates that C. vil data on flight periods in the Dovrefjell montains. losa is better adapted to survive low temperatu Denis (I 98 0 found that imaginal quiescence of res than summer caddisfly species, and in habi egg development in many limnephilids was con tats where frequent freeze-thaw cycles occur, it trolled by the day length the specific larval in may be ecologically advantageous for a species stars experienced. Long day was followed by a to super-cool, at least moderately, to avoid dis direct development, and short days resulted in a ruption of the life cycle by a cold tolerance me quiescence. In the Dovrefjell mountains larvae chanism (Block 1982). are exposed to long days, more than 12 hrs, du A. zonella and C. villosa have a slow growth ring April to October, and direct egg develop and require several years to complete their life ment is to be expected, and was found. cycles. An opposite strategy was demonstrated Crichton (I 97 0 and Crichton and Fisher by the univoltine A. lapponicus. This species in (I 98 0 divided the flight periods of limnephilid habits temporary pools and has a fast growth caddisflies as follows: and completes the maturation of larvae and pu I. An extended flight period probably invol pae within a few weeks (Solem 1983c). A. lappo ving a diapause, from spring through sum nicus is a shredder, and fast growth ofthe larvae mer into autumn; requires reasonably high temperatures and food 2. a shorter flight period, without a diapause, of high quality. The habitats occupied by A. lap in spring and summer, and sometimes ex ponicus are Shallow pools and the water reaches tending into autumn; fairly high temperatures as soon as the ice and 3. a short flight period, without a diapause, in snow have melted. It is also a reasonable as autumn. sumption that the food is ofhigh quality because None of my species is applicable to Crichton's Biirlocher et al. (1978) in temporary pools in I st group. When excluding the spring aspect in southern Ontario reported that detritus in water group no. 2, most of the species, e.g. A. mulieb less pools basins containes more protein than si ris, A. zonella, A. hispida, P. picicornis, L. panto milar detritus in sub-merged areas. dapus and the non-limnephilids belong to group

76 no. 2. Group no. 3 is represented by H. radiatus, Hordaland county (Andersen, T. 1978) show fa H. digitatus and C. villosa. A fourth group that irly similar compositions ofspecies, and indicate is present in the Dovrefjell mountains is the late that the proportion ofthe functional feeding gro winter species, C. sieboldi, which is present in ups revealed at Dovrefjell are representative also April-May. If I divide group 2 into short and ofother montainous areas of South and Central long flight periods, the phenological types are si Norway. Combining the middle and the low al milar also to the distinction made by Malicky pine zones, the scrapers are the dominant group (I 978). here, with the shredders and the predators do The emergence and flight periods of caddisf minant in the sub-alpine birch forest. lies do not necessarily cover the same time Wiggins and Mackay (I978) gave the propor period. Species that live in intermittent streams tions ofthe various functional feeding groups in and temporary pools often have an emergence caddisfly communities in North America (Table period restricted to spring, but have a flight 12). The dominating species in the sub-alpine period that extends into autumn (Crichton 1971, birch forest at Dovrefjell are shredders and pre Svensson 1972, Malicky 1981). The emergence dators, and deviate from the proportions repor and Malaise trap collections in the present study ted by Wiggins and Mackay (I 978). However, had great similarities and was the most common their basic unit is the genus which they regarded pattern in caddisflies in the Dovrefjell mounta as a taxonomic and ecological unit. As I have ins. Only A. muliebris had a delayed peak in Ma used the species as a basis in the description of laise trap captures when compared with that of the functional feeding units this might be a rea emergence traps. son for the differences. Vannote et al. (I980) The sex ratios of P. picicornis was opposite to gave approximate proportions of the functional each other in emergence and Malaise trap collec feeding units for the total invertebrate commu tions, 24.5% females were counted in the emer nity in North American streams, but the Tri gence traps while only 78.6 % females were cap choptera as a group at Dovrefjell differ from tured in the Malaise traps. This suggests that the their scheme also. The proportions found in the swarming or flight prior to mating occurred at present study are different also from the general areas not covered by the Malaise traps above the picture described by Winterbourn et al. (I981) running water. For most species the similarities for New Zealand streams. It is inferred by Van in the sex ratio between the emergence and Ma note et al. (I 980) that their model of the struc laise traps indicate very similar flight habits and ture and function of stream ecosystems is uni flight areas of males and females. However, Ma versal. It is too early yet to say how the total pic laise traps above running water and on the ture will look in Norway, but what has been banks of the Jerosbekken gave a sex ratio of P. shown here is that Norwegian Trichoptera com cingulatus very close to I: I above running wa munities are different from what is assumed to ter while it was about 3 males to I female on the be the structure and function of North Ameri banks. Pre-mating flight of P. cingula/us is car can Trichoptera communities. This implies that ried out in'the bushes on the bank (own obser other groups ofaquatic invertebrates may be dif vations), and a similar feature may be the case ferent from the basic units used by Vannote et also in P. picicornis. al. (I 980), if their general model should be simi The sex ratio of P. conspersa was fairly simi lar in North America and Norway. lar in the emergence and Malaise traps, 21. 8% and 18.2 % females respectively, and resembles sex ratios found earlier, 8.8 % females in light traps by Svensson (I 972), 15.7% females in ACKNOWLEDGEMENTS light traps by Crichton et al. (I 978), and 4.1 % females in light traps by GOthberg (1970). Ho I acknowledge the assistance in collecting mate wever, Tachet (I 967) recorded 57 % females in rial received by 1. Andersen, J. Steinkjer, O. emergence traps and Svensson (1972) 48 % fem Frengen and my wife Thyra Solem. S. Bretten ales in Malaise traps. The different sex ratios re has in various ways given help. A. Lillehammer corded in Malaise traps, may be explained by has given valuable comments to the manuscript. different collecting sites of the traps. I Harder typed the manuscript and Ian Crichton Collections of caddisflies in the mountains of corrected the English. Financial support was gi Saltfjellet, Nordland county (own unpubl. col ven by the Norwegian Research Council for Sci lections), Jotunheirnen, Oppland county (Lille ence the the Humanities, grant nos. D.65.73-10 hammer 1978, own colI.) and Hardangervidda, and D.65.73.032. 77 REFERENCES l..epneva, S.G. 1971. Larvae and pupae of Integripal pia, Trichoptera. Fauna ofthe U.S.s.R. Zool. Inst. Andersen, J. 1984. Biologi og livssyklus tif Chaetopte Acad. Nauk. S.S.S.R. new Ser. 95, 1-560. ryx villosa (Fabricius) (Trichoptera, Limnephili (Trans. Israel Program Sci. Trans. Inc). dae) i og ved Blesbekken og Raubekken, Kongs Li11ehammer, A. 1978. The Trichoptera of 0vre He volt, Oppdal. Unpubl. thesis, Univ. Trondheim. imdalsvatn. Holarct. Ecol. 1, 255-260. Andersen, T. 1974. Caddisflies (Trichoptera) from Lillehammer, A. & Brittain, 1. 1978. 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Trichoptera, 1977. turation genitale des femelles de quelques Lim Junk, The Hague. nephilides. Pp. 57 -66 in Moretti, G.P. (Ed.), Solem, J.O. 1979. A comparison of species diversity Proc. 3rd Int. Symp. Trichoptera. The Hague. indices in Trichoptera communities. Pp. G6thberg, A. 1970. Die Jahresperiodik des Trichop· 255-267 in Grassle, 1.F., Pati!, G.P., Smith, teren imagines in zwei lappHindischer Bilchen. W.K. & Taillie, C. (Eds.) Ecological diversity in Ost. Fish. 23, 118-127. theory and practice. International Co-operative Krebs, C. 1972. Ecology. The experimental analysis of Publishing House, Burtonsville, Maryland. distribution and abundance. 2nd ed. Harper Inter. Solem, J.O. 1981. Overwintering strategies in some Edition. Norwegian caddisflies. Pp. 321-322 in Moretti,

78 G.P. (Ed.) Proc. 3rd Int. Symp. Trichoptera, Series Tachet, H. 1967. Quelques aspects du cycle biologi Entomologica, 20. Junk, The Hague. que de Plectrocnemia conspersa (Curtis 1834) Solem, J.O. 1983a. Identification of the Norwegian (Trichoptera, Polycentropodidae). Annls Limnol. larvae of the genus Potamophylax Wallengren, 3,177-184. 1891 (Trichoptera, Limnephilidae), with data of Tobias, W. 1981. Eine neue Apatania-Art aus Nor life histories, habitats and food in the Kongsvoll wegen (Trichoptera, Limnephilidae). Entomol. area, Dovrefjell mountains, Central Norway. Zeilschr. 91, 62-65. Fauna norv. Ser. B 30, 69-76. Tobias, W. & Tobias, D. 1971. KOcherfliegen und Solem, J.O. 1983b. Temporary pools in the Dovre Steintliegen einiger Gewiisser in S0r-Varanger mountains, Norway, and their fauna of Trichop (Nord-Norwegen) (Trichoptera, Plecoptera). tera. Acta Entomol. Fennica 42, 82-85. Senckenbergiana bioi. 52, 227 - 245. Solem, J.O. 1983c. Larval and pupal descriptions of Ulfstrand, S. 1970. Trichoptera from the River Vin Asynarchus contumax McLachlan, 1880 and A. deliilven in Swedish Lapland. A four-year study lapponicus Zetterstedt, 1840, with notes on biono based mainly on the use of light-traps. Entomol. mics in Centr. Norway (Trichoptera: Limnephili Tidsskr. 91,46-63. dae). Ent. scand. 14,439-451. Vannote, R.L., Minshall, G.W., Cummins, K.W., Se Solem, J.O. 1983d. Identification of Norwegian ge dell, J.R. & Cushing, C.E. 1980. The river conti nera of limnephilid larvae with single-filament nuum concept. Can. J. Fish., Aquat. Sci. 37, gills (Trichoptera: Limnephilidae). Ent. scand. 14, 130-137. 457-461. Wallace, J.B. & Merritt, R.W. 1980. Filter-feeding Solem, J.O. 1984. Adult behaviour of North Euro ecology of aquatic insects. Ann Rev. Ent. 25. pean caddistlies. Pp. 375 - 382 in Morse, J.c. 103-132. (Ed.) Proc. 4th Int. Symp. Trichoptera, Series En Wiggins, G.B. 1973. A contribution to the biology of tomologica, 30. The Hague. caddistlies (Trichoptera) in temporary pools. Life Solem, J.O. 1985. Norwegian Apatania Kolenati (Tri Sci. Contr., R. Onto Mus. 88, 1-28. choptera: Limnephilidae): identification of larvae Wiggins, G.B. & Mackay, R.J. 1978. Some relations and aspects of their biology in a high-altitude hips between systematics and trophic ecology in zone. Ent. scand. 16, in press. nearctic aquatic insects, with special reference to Solem, J.O., Sendstad, E., Bergvik, T. & Hegstad, A. Trichoptera. Ecology 59, 1211-1220. 1977. New record of Apatania zonella Zett. (Tri Winterbourn, M.J., Rounick, J.S. & Corvie, B. 1981. choptera, Limnephilidae) from Svalbard. Norw. J. Are New Zealand stream ecosystems really difTe Ent. 24, 85. rent'? N. Zealand J. Mar. Freshw. Research 15, Svensson, B.O. & Tjeder, B. 1975. Check-list of the 321-328. Trichoptera of North-Western Europe. Entomol. scand. 6, 261-274. Received I Febr. 1985 Svensson, B.W. 1972. Flight periods, ovarian matu ration, and mating in Trichoptera at a South Swe dish stream. Oikos 23, 370-383.

79 Female sex pheromones in Rhyacophila nubila (Zetterstedt) (Trichoptera, Rhyacophilidae) and arrival pattern to sticky traps

JOHN O. SOLEM

Solem, 1.0. 1985. Female sex pheromones in Rhyacophila nubila (Zetterstedt) (Trichoptera, RhyacophiJidae) and arrival pattern to sticky traps. Fauna norv. Ser. B. 32. 80-82.

Using live caged females and extracts offemales ofthe caddislly Rhyacophila nubila (Zetter stedt), I have shown female sex pheromones to be present. Live caged males did not attract females. The pheromones are produced on sternite 4; they are present in very young pupae, and probably in old prepupae also. The peak in the arrival pattern of adults at the sticky traps was in early night. lohn O. Solem, University of Trondheim, The Museum, ErJing Skakkesgt. 47A, N-7000 Trondheim, Norway. INTRODUCTION Within the insect order Trichoptera we know one to five days old, were placed ~eparately in of two behavioural patterns used when sexes cylindrical screened cages (30 x 45 mm). Two meet for reproduction. One pattern is that the females which were reared together with several male approaches the female and this is probab individuals, and had most likely mated, were ly the most wide-spread and ancient pattern. It also used. The cages were placed in square has been reported from several families (Kel sticky traps measuring 20 x 30 cm. ner-Pillault 1975, Solem 1976, 1984, Wood Various parts ofthe body offemales were pla and Resh 1984). The second pattern is that the ced in a glass vial with 0.5 to I ml of dichloro females fly into a swarm of males, and make methane. After 1.5 hrs extraction the tissue physical contact to facilitate sexual recognition. were removed, and the extracts were infused to This second pattern has been reported in fam. rubber septa (F-138 Septa from Supelco Inc.). Hydropsychidae (Fremling 1960) and fam. Lep The extracts were made just before darkness. In toceridae (Hickin 1953, Solem 1978, 1984, order to isolate the segment that produced the Tozer et al. 1981). It seems reasonable to pheromones, the abdomens were dissected and assume that the two patterns have different the different parts tested against each other for attraction mechanisms. The attraction mecha male attraction; e.g. segments one to four were nism in leptocerids and hydropsychids is un tested against the rest ofthe abdomen, the sterni known, but recently Wood & Resh (984) de tes of segments one to four were tested against monstrated that female sex pheromones are the tergits of segments one to four, the sternites involved in the attraction of males in the seri ofsegments one and two were tested against the costomatid Gumaga griseola McLachlan. sternites ofsegments three and four, the sternite This paper reports the results of field experi of segment three were tested against sternite of ments concerning sexual communication in segment four, and at last, the sternite ofsegment Rhyacophila nubila (Zetterstedt) (Rhyacophili four were tested against the rest of the body. dae). Extracts of females collected in light traps, mature pupae, young pupae, and prepupae were also tested. MATERIAL AND METHODS The traps were placed in the field before dark Field experiments were carried out at the Driva ness. In early August they were checked regu River (880 m a.s.l.) at Kongsvoll Biological Sta larly during the night. The number ofspecimens tion in the Dovrefjell mountains, Central Nor caught was counted in order to get information way, during the summers 1983 and 1984. Pu about the arrival pattern of adults. In late Au pae were collected and reared in separate cham gust the traps were checked only one time du bers until adult stage. Virgin males and females, ring a night.

80 Fauna norv. Ser. B. 32: 80 - 82. Oslo 1985. Table I. Pheromone tests of Rhyacophila nubila

Average number of males Test catched per No. of no. trap/day traps Controls 0.7 10 Live females 10.66 13 p <0.01 Live males 0 3 2 Females, extract (2 inds) Head + thorax 1.0 3 Abdomen 24.33 3 p <0.001 Controls 0.6 3 3 Females from light traps, extracts Abdomen (3 inds) 32 2 p <0.001 Controls 0.75 4 4 Female pupae (mature) Extracts (5 inds) 9 3 p <0.01 Controls 0.75 4 5 Female pupae (young) Extracts (3 inds) 20.25 4 p <0.001 Controls 2.4 8 6 Pre-pupae Extracts (3 inds) 13 2 p <0.05 Controls 4 4 Extracts (3 inds) 2 Controls 1.3 3 7 4th sternite of female abdomen Extracts (3 inds) 38.8 5 p <0.001 Female minus 4th sternite Extracts (3 inds) 1.75 4 Controls 1.3 3

Rhyacophila nublla ....-- 21 to 24 Aug. 1984 1 to 7 Aug. 1984 160

20 120

80 lfJ ...Q) .0 Hrs. 23 02 04 E 40 ~ Fig. l. Numbers of Rhyacophila nubi/a, males and females, which arrived at sticky traps from 2100 to 0400 hours during I to 7 August 1984, and 2100 to 0800 hours during 21 to 24 August 1984. Hrs. 23 08

81 RESULTS uld be identified as glands for releasing phero Data from the field experiments are presented in mones. Table I, and they show that: I) Live virgin fem Solem (J 984) reported that R. nubila congre ales attracted males; 2) Live males did not attract gated and flew in bushes along the banks of females; 3) Extracts of abdomen of virgin fema streams and rivers, but detailed features in the les attracted males, while head and thorax were flight pattern were not detected. This flight is not attractive; 4) Extracts ofabdomen of females very likely males searching for females, because which were caught in light traps, attracted the females rarely fly on the bank (own unpubl. males; 5) Sternite 4 was most attractive to males; data). The experiments carried out on R. nubila 6) Young pupae were also attractive; 7) One trap have focused on long distance pheromones. It is with extracts of prepupae attracted a significant of paramount importance for the interpretation higher number of males than the controls. Ho of the flight behaviour in caddisflies to know if wever, three other traps baited with prepupal such pheromones are present or not. extracts did not differ from the controls. The highest number of R. nubila males were caught in early night, just after darkness (Fig. I). ACKNOWLEDGEMENTS This arrival pattern corresponds to that of light I acknowledge the help received from R. Schmid traps reported by Gothberg (1978). and H. Mustaparta.

REFERENCES DISCUSSION Birch, M.C. 1974. Pheromones. Nort!) Holland, Am The attraction of males to females and to abdo sterdam. minal extracts of females indicate that female Fremling, C.B. 1960. Biology and possible control of sex pheromones are involved in the sexual com nuisance caddisflies of the Upper Mississippi Ri munication in R. nubila. The presence of female ver. Tech. Res. Bull. Iowa State Univ 483 sex pheromones using extracts has been demon 856-879. ., strated only once before in the order Trichop GOthberg, A. 1978. Diel flight activity of some Tri tera, in Gumaga griseola McLachlan (Sericosto choptera in Northern Sweden. In: Crichton, I.M. matidae) (Wood and Resh 1981). R. nubila and (Ed.) Proc. 2nd Int. Symp. Trichoptera, /977. Junk, The Hague. p. 147. G. griseola belong to two different groups of Hickin, N.E. 1953. Courtship ofa caddislly, Mystaci caddisflies. des nigra L. Entomologists Mon. Mag. 89, 71. However, Kelner-Pillault (J 975) reported that Kelner- PilIault, S. 1975. Attirance sexuelle chez un caged females attracted conspecific males in Trichoptere: Enoicyla pusi/la Burmeister. Bull. Enoicyla pusilla Burm., and similar phenome Soc Entomol. Fr., 80, 252-257. non was reported in Phryganea bipunctata (Ret Solem, J.O. 1976. Studies on the behaviour of adults zius) by Solem (1976). Even though, the two last of Phryganea bipunctata and Agrypnia obsoleta observations are not as conclusive as using fem (Trichoptera). Norw. J. Ent. 23, 23-28. ale extracts, they do indicate that female sex Solem, J.O. 1978. Swarming and habitat segregation in the family Leptoceridae (TrichopteraJ Norw. J. pheromones are present in many different gro Ent. 25, 145-148. ups ofcaddisflies, and may be they will show to Solem, J.O. 1984. Adult behaviour of North Euro be as common as in the sister order Lepidoptera. pean caddisllies. In Morse, l.C. (Ed.) Proc. 4th Int The attraction of males to one trap baited with Symp. Trichoptera. Series Entomologica 30. Junk, extracts of prepupae indicates that pheromones The Hague. pp. 375-382. might be produced at the prepupal stage. It is Tozer, W.E., Resh, V.H. & Solem, J.O. 1981. Biono possible that these prepupae were older than mics and adult behaviour of a lentic caddislly, those used in the three other, which were not at Nectopsyche albida (Walker). Am. MidI. Nat. /06, tractive. Testing sternite 4 against the rest of the 133-144. Wood, J.R. & Resh, V.H. 1984. A demonstration of body is a strong indication that pheromones are sex pheromones in caddisllies (Trichoptera). J. produced on sternite 4. However, glands in this Chem. Ecol. 10,171-175. segment have not been detected. SEM photos of the sternite did not show any structure that co- Received I Febr. 1985

82 The geographical range extension ofHeterhelus scutellaris (Heer 1841) (Coleoptera, Nitidulidae) and its host plant Sambucus racemosa (L.) (Caprifoliaceae) in Scandinavia

PREBEN OTIESEN AND TORSTEIN KVAMME

Ottesen, P. & Kvamme, T. 1985. The geographical range extension of Heterhelus scutellaris (Heer 1841) (Coleoptera, Nitidulidae) and its host plant Sambucus racemosa (L.) (Caprifolia ceae) in Scandinavia. Fauna norv. Ser. B. 32,83-88.

In Scandinavia Heterhelus scutellaris (Heer 184 I) was recorded for the first time in Finland 1901, Denmark 1916, Sweden 1947 and Norway 1979. At present the species is exceedingly common in Southern Scandinavia, and is still spreading. Two independent dispersal routes have been identified: across the Gulf of Finland to Finland, and from Germany to Denmark. The latter population has spread to Sweden and further to Norway. This natural dispersal has followed the human introduction and cultivation of its host plant, Sambucus racemosa (L.), which 100 to 150 years ago rapidly became naturalized through seed dispersal by birds. In Europe both species today have disjunct distribution patterns: the new Scandinavian one, and the original one at higher elevations in the central and southern parts ofCentral Europe. The warm lowlands in between apparently act as a barrier. In Scandinavia the beetle has to day reached 300 km south ofthe northern distribution limit of fertile S. racemosa, the latter being found at 65°N.

Preben Ottesen, Department of Biology, Division of Zoology, University of Oslo. P.O.Box 1050, Blindern, N-0316 Oslo 3, Norway. Torstein Kvamme, Norwegian Forest Research Institute, P.O.Box 61, N-1432 As-NLH, Norway.

INTRODUCTION Geographical range extensions ofanimal species the beetle, compiled published and unpublished are frequently reported, but the reasons underly records up to 1983. ing such ex~nsions are rarely known (Krebs 1972). The best understood cases are found when monophagous species have spread to BIOLOGY areas where their host plants have been introdu H. scutellaris is a 2.0 to 2.5 mm long, yellow ced. However, few good examples are available reddish beetle which belongs to the subfamily (Lattin 1967). Kateretinae of the Nitidulidae family (Fig. I). It The present paper will document the natural is readily ,!;stinguished from related species in dispersal of the Nitidulid beetle Heterhelus scu having two sep.:ments only (not three) in the an tellaris (Heer 1841) in Scandinavia. This disper tennal club (Hansen 1950). Its life history is clo sal has clearly been possible due to the introduc sely associated with the bush S. racemosa on tion and ensuing naturalization of its host plant which larvae and imago feed on pollen. Other Sambucus racemosa (L.). Although Scandina parts of the flowers may be eaten as well. Occa vian coleopterologists have recorded catches of sionally imago may visit other flowering trees, the beetle for more than eighty years and bota like Sorbus aucuparia L. Pupation takes place in nists have followed the bush for more than 150 or on the ground below fallen flowers (Rupert years, all information on the two species con sberger 1872). Its annual period ofactivity coin cerns single records or observations within limi cides with the flowering of S. racemosa, which ted geographical areas. To obtain a comprehen is in the spring, usually in Mayor June. It is of sive picture oftheir biology, range extension and ten seen flying around the bushes and is usually current distribution, we have, with emphasis on found in large numbers in the flowerheads.

Fauna norv. Ser. B. 32, 83-88 Oslo 1985. 83 LIST OF RECORDS quickly established itself in Skane. On 25 May 1950 it was registered to be common on Sorbus The district abbreviations follow Lindroth aucuparia and S. racemosa in Sk, Sovde (Palm (1960), but the faunal divisions for Norway ag 1950, and Israelson (I 956) found it in several ree with 0kland (I 981). EIS-numbers for Nor localities in the Hiissleholm area, Sk on every way refer to the European Invertebrate Survey, bush he investigated, even isolated ones. In 1957 Norwegian grid numbers (0kland 1976). two specimens were caught, at least one on S. aucuparia, in Vg, GOteborg (ArvalO, and on 29 Finland April 1961 it was found to be numerous on S. The first record of H. scutellaris in Scandinavia racemosa in the same area (Hagstrom 1962). It is from Finland. In 190 I several specimens were was also recorded in Srn by Dahlgren (Klefbeck found on S. racemosa close to Helsinki (Sahl 1962). On 6 June 1962 it was found in Hall, Fja berg 1902). We have not been able to find any rasbracka (Lundberg 1963) and the same year in other published records, but Olof Bistrom has BI, Listerby by N. Gyllensvard (Lundberg 1968). kindly sent us information on the specimens in Early in the seventies the species appeared in se the Helsinki Zoological Museum. The years of verallocalities ofCentral Sweden. The first three first capture in the various faunal provinces of records were all from Dlr, which is remarkably Finland, judged from this material, are: N: 190 I, far north in relation to the previous Swedish Ik: not dated, but probably collected early in this catches: Dlr, Borlange, 26 May 1971 (G. Sjodin); century, Ta: 1902, Ab: 1914, Sa: 1932, Kl and Dlr, Orsa, 2 June 1971 (A. Petrelius); and Dlr, Ka: 1938, Tb: 193? (not specified), St: 1952, and Sater, 17 May 1972 (B. Ehnstrom). All subsequ Sb: 1956. It is also found in Kon (Lindroth ent recordings are further south: Vrm, Eje, 25 1960). In several of these provinces the species May 1972 (B. Ehnstrom); Og, Omberg, 6 June has later been rerecorded (0. Bistrom, pers. 1973 (A. Lindelov); Upl, Lunda, 15 Sept. 1973 commJ (A. Anderberg); Sdm, Vardinge, 4 April 1974 (c. Holmquist); and 01, 1983 (J.R. Bergwall). In Denmark several ofthe districts mentioned above the spe The first Danish specimen of H. scutellaris was cies has later been rerecorded, often in large caught on 8 June 1916 in Boiler Skov in the ne numbers (S. Lundberg, pers. commJ ighbourhood of Horsens, Jylland (H0eg 1919). The next record was on 20 May 1920 at N0r Norway holm close to Esbjerg, Jutland, where three spe cimens were beaten down from flowering Sor The first Norwegian specimen was found on 12 bus aucuparia (Larsen 192 3- 24). The third re July 1979 at AK: Asker, Semsvannet (EIS:28) cord was from Svendborg, Fyn, where the spe (P.OJ The next year, 1980, it was found in large numbers on S. racemosa at AK: Brerum, Jar cies was found in large numbers on S. racemosa (EIS: 28), 17 May (A. Fjellberg); eleven speci (West 1936-37) (the bush was published as S. mens in AK: Frogn, Dmbak (EIS:28), 20 May ebulus, but corrected by West (I 947». The date (R. FardaO; and several specimens in YE: Hor was 19 May, but the year was not given, al ten (EIS: 19), 27 May (T.KJ In 1981 it was though it probably was from the early thirties. found in large numbers on every single bush of West (I 940 - 4I) concluded that the beetle was S. racemosa that was investigated in the Oslo «very rare», although one additional record was area (EIS:28), and it was also found in 0: Hal given in the addendum: Fmslev Plantage, Zea den (EIS:20), 27 May and AK: Ullensaker, Hers land, 2 June. However, some years later West joon (EIS:37), 26 May (P.OJ Between 26 May (I 947) gave eight additional records from var and 3 June 1982 it was found on every single ious parts ofthe country: Oens, liylke, Slukefter bush of S. racemosa that was investigated in co (NW Odense), Boserup, Nybro Skov, Fures0 astal districts of Southern Norway: 0: Fredrik parken, Gelsskov, 0rholm - and concluded stad (EIS: 20); B0: 0vre Eiker, Fiskum (EIS:27); that the species had spread considerably the last B0: Kongsberg, Saggrenda (EIS: 27); B0: Kongs years. This observation was confirmed by Han berg city (EIS:27); TEY: Skien (EIS: 18); TEY: sen (I952). Bamble, Valle (EIS: 10; and AAY: Arendal (EIS:6) (T.KJ On II July 1982 a search was Sweden made for the species in BV: AI (EIS:43). Bet The first Swedish specimen was caught at Sk, ween the railway station and Al centre 705 flo Bokeberg on 27 May 1947 (Leiler 1947). It werheads on a total of 31 bushes ofS. racemosa

84 were examined. Only five specimens were mentioned above, is a typical habitat for the spe found, although S. racemosa was abundant in cies in Southern Germany. the area and all the way down the Hallingdal The rapid dispersal in Scandinavia is most lik valley southwards toward the coast (P.O.). ely due to two factors: man and birds. Lagerberg et al. (1957) wrote that the bush was frequently used to camouflage «certain necessary small DISCUSSION buildings» at railway stations all over Norway! Also in Sweden the bush was frequently planted The range extension ofS. racemosa around railway stations and other railway buil The range extension of H. scutellaris in Scandi dings (Almquist 1928). Concerning birds, it has navia is undoubtedly connected with the range long been observed that a number of species extension of its host plant Sambucus racemosa, feed on its red berries. Holmboe (I 900) reported which is found throughout the Holarctic region. that the berries were successfully used as bait in Prior to the invasion of Scandinavia, the nort thrush (Turdus sp.) traps, and Schuster (I930) hern range of this bush in Europe stretched listed 36 species ofbirds which have been obser from Belgium and Central Germany to Lithua ved to feed on the berries of S. racemosa. Mei nia, and in the south from the Pyrenees to dell (1937) found seeds ofS. racemosa in the sto Southern Bulgaria (Tutin et al. 1976). As in machs of fieldfares (Turdus pilaris L.) and noted Scandinavia it has become naturalized in the that they were all undamaged. It is therefore USSR from Lithuania to Lake Ladoga, although reasonable to conclude that the main factor in scattered from Lake Peipus (between the two ar the naturalization of the species is birds. How rows on Fig. I) and eastwards (Hulten 197 I). ever, vegetative reproduction is quite successful, The bush probably prefers cold climates. It does and may locally produce dense stands in cold not occur naturally in the warm northern low areas where fertile specimens are absent (Con lands of Central Europe, including Great Bri rad & McDonough 1972). tain, although it was probably present in the lat ter country in earlier interglacials (Godwin 1975). In Southern Germany it is considered to The range extension ofH. scutellaris be a characteristic species of the phytosociologi H. scutellaris is found in Europe to East Siberia cal class Epilobietea angustifolii, alliance Sam (Reitter 1919), but not, like its host plant, in the. buco-Salicion capreae, which typically occurs Nearctic region. Its range in Europe coincides on fresh, loose, humic soils with a good nutrient with that of S. racemosa (Freude et al. 1967) supply, especially on old forest felling areas, and (Fig. I). Both organisms are thus rare in the usually in submontane and montane regions (0 warm lowland parts of Northern Central Eu berdorfer 1973). rope. However, once this climatic barrier was More than 150 years ago the bush was intro crossed, the two species have obviously met ex duced to Standinavia and cultivated for orna cellent conditions in the cooler areas further mentation (Lagerberg et al. 1957). With the aid north. ofman, the warm Central European barrier was Fig. I shows the range extension ofH. scutel thus crossed. In 1837 it was first observed to laris in Scandinavia. It must be emphasized that spread naturally in the Stockholm area (Hylan the 10-year distribution fronts are approxima der 1971), and around 1870 in the Oslo and Ber tions only, since the number of observations is gen areas (Blytt 1874, Holmboe 1941). The follo few, and since the years offirst captures are pro wing natural dispersal was fast, and within a bably not identical with the years of coloniza few decades the species was fairly common in tion. However, several trends are apparent and Sweden (Almquist 1929). Schtibeler (I888) re will be discussed below. ported that the species thrived well in Norway, H. scutellaris has obviously entered Scandina even close to the timberline, and in severalloca via via two independent routes: to Finland lities it spread like a weed. Today the bush is across the Gulf of Finland or from the Lenin common over the whole of Southern Scandina grad area (Fig. I, two arrows), and to Denmark via, especially in areas with human settlement. from Schleswig-Holstein, Germany (Fig. I, ar Although its northern distribution limit has pro row). The distance across the Gulf of Finland bably been reached (Fig. I), it still seems to be seems short, but in the previous century S. race expanding within the present range. In recent mosa was probably far more uncommon along years it has become increasingly common on old the USSR coast than it is today (see discussion forest felling areas (Lundqvist 1983), which, as under S. racemosa). In Finland H. scutellaris ra-

85 Fig. I. The range extension of H. scute//aris in Scan bars (rT""T"ll. The distribution of S. racemosa along dinavia shown by approximate IO-year distribution the USSR coast of the Baltic is, according to Hulten frol1ts. In some districts of Central Sweden and (J 971), a result of recent cultivation, and the bush is South-Eastern Norway the years of first captures are still scattered from Lake Peipus (between the two ar indicated. Arrows (~) indicate plausible routes of in rows) and eastwards. vasion. Shaded areas ( ~ ) show the distribution of S. racemosa in Northern Europe according to the map by Hulten (1971), and in Central Europe, where pidly spread towards the north, apparently with limited by a solid line (--), according to the map by a speed of 5- 10 km a year, until the thirties Waldenburg (1934), while the remaining southern when expansion seems to have ceased around areas are drawn approximately according to text in 62.5°N. However, this should not be taken for formation from Tutin et al. (1976). The bush is not a native of Great Britain. granted until northern Finland is more closely According to R. Elven (pers. commJ, the shaded investigated. In Denmark the species has expan areas show the distribution of fertile S. racemosa, ded at approximately the same speed as in Fin while vegetatively reproducing specimens in Norway land, 6-10 km a year. In the mid-forties it may be found north to the solid line with vertical spread from Denmark to Sweden and continued 86 to disperse north and east. The speed ofdispersal even further north if flowering bushes of S. ra seems to have increased during the last decades. cemosa are found in spring. The species should During the seventies several specimens were also be expected to colonize the coastal districts found in Central Sweden, and the species was of Western Norway, where S. racemosa is com obviously well established in Dlr as early as mon. However, the high mountains may repre 1971. The dispersal from Southern Sweden may sent a geographical barrier that will retard dis thus have been faster than the distribution fronts persal. indicate. Similarly, that the beetle was common in all South-Eastern Norway when it was first actively searched for, strongly indicates that it ACKNOWLEDGEMENTS had been present for several years prior to its We would like to thank Stig Lundberg, Lulea, discovery. Another explanation of the apparent and Olof Bistrom, University of Helsinki, for rapid expansion in Sweden may be that the Fin kind permission to use their unpublished Swe nish population has spread across the Gulf of dish and Finnish material respectively, Sverre Bothnia via the Aland archipelago (Fig. 1, arrow L0kken, University ofOslo, for extensive infor with ?). However, no records ofthe beetle from mation on botanical literature, Reidar Elven, these islands exist, although this may be due to Troms0 Museum, for information on S. race insufficient searching. It must be once more re mosa in Northern Norway, and Robert Vinter, membered that observations are too few and ca University of Oslo, for improving the language. usal to allow any exact interpretation. Human introduction of H. scutellaris should not be completely ruled out. If young seedlings of S. racemosa were introduced, the beetle, REFERENCES which pupates and hibernates below the bushes, A1mquist, E. 1929. Upp1ands vegetation och flora. may have followed in the soil. According to Acta phytogeogr. suec. 1, 622 pp. Hansell (1969), new bushes even grow from pie Blytt, M.N. 1874. Norgesjlora. Del 2: Med histand af ces of root which have been thrown away. Ho Professor M.N. Blytts efterladte Optegnelser og wever, since the species is a good flier, and since Samlinger, afAxel Blytt. Kongelige Norske Vit. it has entered the southern areas first, it most lik Selsk., Christiania (Oslo). ely has dispersed naturally with the wind. Conrad, P.W., & McDonough, W.T. 1972. Growth For permanent colonization to be successful, and reproduction of red elderberry on subalpine the chance offinding a favourable site for repro range1and in Utah. Northwest Sri. 46, 140-148. duction must be greater than the risk of extinc Freude, H., Harde, K.W., & Lohse, G.A. 1967. Die H. Kafer Mitteleuropas. 7. Clavicornia. Goecke & tion. The establishment of scutellaris in Scan Evers. Krefeld. dinavia was probably first possible when S. ra Godwin, H. 1975. The history of the British jlora. cemosa had reached a certain critical density le 2nd. ed. Cambridge Univ. Press. Cambridge. vel, which may explain the time lag between the Hagstrom, T. 1962. Coleoptero10giska medde1anden. initial naturalization of the plant and the arrival Opusc. ent. 27, 59-60. of the beetle. Thus, the results above indicate Hansell, O.A. 1969. Bdrbocken. P.A. Norstedt & Sa that such a level was reached somewhere bet ner. Stockholm. ween 50 and 100 years after the naturalization Hansen, V. 1950. Biller 13. Clavicornia (I). Danm. of the bush had started. Fauna 55, 278 pp. - 1952. JEndringer i den danske billefauna i nyere At present the Scandinavian distribution ofH. tid. Ent. Meddr 26, 438-448. scutellaris has reached about 300 km south of - 1964. Fortegnelse over Danmarks biller (Coleop the northern distribution limit of S. racemosa tera). Ent. Meddr 33, 1-506. (Fig. I). The latter limit, which is taken from H0eg, N. 1919. Nogle meddelelser om fund af sjreld Hulten (1971), seems to apply to fertile bushes. nere og enkelte for den danske fauna nye bille Non-reproductive bushes may be found up to arter i Jylland, specielt Horsensegnen. Ent. Meddr 69°N latitude (Reidar Elven, pers.comm.), but 13. 23-29. these are of course useless to the pollen-feeding Holmboe, J. 1900. Notizen iiber die endozoische beetle. Samenverbreitung der Vogel. Nyt Mag. Naturvid. 38,305-320. It will be interesting to see how far the beetle - 194 I. Gratis mat av ville planter. J.W. Cappelen. will extend its range, i.e. whether the climatic Oslo. tolerance ofthe beetle will allow it to invade the Hulten, E. 1971. Atlas over viixternas utbredning i entire range of its host plant or not. The species Norden. Generalstabens 1itografiska anstalts for should be looked for up to 65°N latitude, or lag. Stockholm. 87 Hylander, N. 1971. Forsta Iitteraturuppgift fOr Sveri 0kland, J. 1976. Utbredelsen av noen ferskvanns ges vildviixande kiirlviixter jiimte uppgifter om muslinger i Norge, og litt om European Inverte fOrsta svenska fynd. Suppl. Svensk bot. Tidskr. 64 brate Survey. Fauna, Oslo 29, 29-40. (1970), 332 pp. 0kland, K.A. 1981. Inndeling av Norge ill bruk ved Israelson, G. 1956. Skalbaggar fran Hiissleholmstrak biogeografiske oppgaver - et revidert Strand-sy ten VI. Natur i G6inge 1-2,7-13. stem. Fauna, Oslo 34, 167 -178. KIefbeck, E. 1962. Catalogus Insectorum Sueciae. Palm, T. 1951. Anteckningar om svenska skalbaggar. XVI. Coleoptera 1960. Nigra fOrklaringar och til VI. Ent. Tidskr. 72. 39-53. liigg. Opusc. ent. 27, 153-174. Reitter, E. 1919. Bestimmungs-Tabelle der europai Krebs, C.l. 1972. Ecology. The experimental analysis schen Coleopterenfamilien: Nitidulidae und Bytu of distribution and abundance. Harper & Row. ridae aus Europa und den angrenzenden Liindern. N.Y. Verh. nalurf Ver. Briinn. 56, 1-104. Lagerberg, T., Holmboe, J., & Nordhagen, R. 1957. Rupertsberger, M. 1872. Beitrage zur Lebensge Vdre ville planter. VI. Johan Grundt Tanum. schichte der Kiifer. Verh. zool.-bot. Ges. Wien Oslo. 1872,7-26. Larsen, F. 1923-24. Fund af danske biller, isrer i Sahlberg, l. 1902. Heterhelus scutellaris Heer. Esbjergegnen. Ent. Meddr 14,228-235. Meddl. Soc. Fauna Flora Fennica 28, 48-49 N Lattin, G. de 1967. Grundriss der Zoogeographie. Schtibeler, F.e. 1888. Viridarium Norvegicum. Nor Gustav Fischer Verlag. Stuttgart. ges Vrextrige. 2. Universitetsprogram. Christiania Leiler, T.E. 1947. Bidrag till kiinnedomen om sven (Oslo). ska coleopterers utbredning och levnadsiitt. Schuster, L. 1930. Ueber die Beerennahrung der Vo Opusc. Ent. 12, 158. gel. J. Om., Lpz. 78,273-301. Lindroth, C.H. (ed.) 1960. Catalogus Coleopterorum Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, Fennoscandiae et Daniae. Entomological society. D.M., Valentine, D.H., Walters, S.M., & Webb, Lund. D.A. 1976. Flora Europaea. 4. Plahtaginaceae 10 Lundberg, S. 1963. Bidrag till kiinnedomen om sven Compositae (and Rubiaceae). Cambridge Univ. ska coleoptera. 6. Ent. Tidskr. 84, 119 -124. Press. Cambridge. - 1968. Catalogus Insectorum Sueciae XVIII. Cole WaIdenburg, I. 1934. Die floristische Stellung der optera 1960. Riittelser och tilliigg. Opusc. Ent. 33, Mark Brandenburg. Verh. bOI. Ver. Prov. 197-218. Branden. 75, 1-80. Lundqvist, R. 1983. Mycket pi gang pi hyggena. West, A. 1936-37. Tillreg og rettelser til fortegnel Svensk bot. Tidskr. 77, I. serne over de danske Coleoptera Ill. Ent. Meddr Meidell, O. 1937. Unders0kelse av mageinnhold hos 19, 452-492. gratrost (Turdus pilaris L.) Nyll Mag. Naturvid. - 1940-41. Fortegnelse over Danmarks biller, 76, 163 - 236. deres udbredelse i Danmark, forekomststeder og Oberdorfer, E. 1973. Die Gliederung der Epiobietea tider, biologi mm. Enl. Meddr 21, 1-664. anguslifoiii - Gesellschaften am Beispiel Stidde - 1947. TiIlreg til «Fortegnelse over Danmarks bil utscher Vegetationsaufnahmen. Acta bot. hung. ler, deres udbredelse i Danmark, forekomststeder 19,235-253. og -tider, biologi mm.» Ent. Meddr 25,3-141.

Received 11 Febr. 1985

88 A check-list of1Norwegian blow-flies (Dipt., Calliphoridae)

KNUT ROGNES

Rognes, K. 1985. A check-list of Norwegian blow-flies (Dipt., Calliphoridae). Fauna norv. Ser. B. 32, 89-93.

Calliphorid material from Norway in the museum collections of the Universities in Bergen, Oslo, Trondheim and Troms0 and in my own collection has been revised and a check-list of the 39 species known at present from Norway is presented. Pollenia obscura Bigot, 1888, preoccupied in Pollenia by Musca obscura Fabricius, 1794, is redescribed and removed {rom the synonymy of Pollenia rudis (Fabricius, 1794); pseudorudis nom.nov. is proposed as a replacement name.

Knut Rognes, Stavanger lrererh0gskole, Postboks 2521 Ullandhaug, N-4001 Stavanger, Norway.

The check-list presented below, which replaces wegian calliphorid fauna. The nomenclature previously published lists (Siebke 1877, Sch0yen adopted is based on Zumpt (956), Peus (960) 1889, Bidenkap 1892, 1901, Ringdahl I944a, and Schumann 0964, 1973a, 1973b, 1974). Sy 1944b, 1952) is based upon a revision of the nonyms have mostly been taken from Aubertin blow-fly material from Norway in the museum (933), Ringdahl (945), Zumpt (956), Peus collections ofthe Universities in Bergen (mainly (960), Schumann 0964, 1973b, 1974), Pont O. Bidenkap, L. Greve, A. L0ken, N. Knaben, 098I) and Rognes 0983b, 1985 and in press). T.R. Nielsen leg.), Oslo (mainly 1. Knaben, Generic synonyms have been cited when the 1.H.S. Siebke leg.), TromS0 (mainly T. Soot type-species is on the list (as valid name or syno Ryen leg.) and Trondheim (DKNVS-Museet) nym), specific synonyms when based on type • (mainly 1.0. Solem leg.)(about 1820 specimens) specimens from Scandinavian type-localities, and in my own collection (about 2500 speci when being the basis for cited genus names or mens). In addition a few small private collec when commonly met with in literature dealing tions have been examined (B. Sagvolden, Rollag, with the Scandinavian fauna. J.H. Simonsen, Oslo). I have only seen parts of Many ofthe currently used names are unfor the V. Storm eolleetion in Trondheim revised by tunately preoccupied and name-changes are to Ringdahl 0944a). I have examined specimens be expected in the future. I have mostly held on • from Norway of all the species listed. Ringdahl to established names, except in a few cases. Neit o944b, 1952) listed 18 species from Norway. her have I made any changes as to the currently Ard6 (1957) added I and Rognes (1980, 1982, recognised generic limits, even though a few 1983a, 1985) 9 bringing the total to 28 species. changes appear quite justified and necessary. The present revision lists 39 species from Nor These are indicated in the annotations at the end way. In comparison it may be noted that 20 spe of the paper. cies have been published from Denmark (Lund In the list generic and specific synonyms are beck 1927), 34 from Finland (Hackman 1980), indented. Superscript numbers refer to the num 32 from Sweden (Ringdahl 1952) and 33 from bered annotations. the British Isles (Pont 1975). Note that Helieobo sea palpalis, treated as a tachinid or sarcophagid by the cited authors, is not included in these Family Calliphoridae numbers. Four subfamilies are represented in the Nor- Subfamily Calliphorinae Bellardia Robineau-Desvoidy, 1830 I) Paper no. 2 in a series based on investigations fi agilis (Meigen, 1826) I nanced by a grant «(vikarstipend») from Stavanger ampleetens (Pandelle, 1896) lrererh0gskole 1982-1983. pusilla (Meigen, 1826) 2

Fauna nory. Ser. B, 32,89-93. Oslo 1985. 89 .~

aculeata (Pandelle, 1896) retrocurva (Pandelle, 1896) zetterstedti (Enderlein, 1933) Pseudonesia Villeneuve, 1920 3 stricta (Villeneuve, 1926) puberula (Zetterstedt, 1838) unxia (Walker, 1849) pubicornis (Zetterstedt, 1838) pandia (Walker, 1849) dolens (Zetterstedt, 1838) biseta (Kramer, 1917) Calliphora Robineau-Desvoidy, 1830 Acrophaga Brauer & Bergenstamm, 1891 Abonesia VilIeneuve, 1927 Subfamily HeJicoboscinae 4 Stobbeola Enderlein, 1933 Helicobosca Bezzi, 1906 4 genarum (Zettersted, 1838) Theria Robineau-Desvoidy, 1830, preocc. laticornis (Zettersted, 1838) palpalis (Robineau-Desvoidy, 1830) alpina (Zetterstedt, 1838) distinguenda Villeneuve, 1924 loewi Enderlein, 1903 germanorum (VilIeneuve, 1907) stelviana (Brauer & Bergenstamm, 1891) norwegica (Enderlein, 1933) alpina: authors, not Zetterstedt (misi Subfamily Phormiinae dentification) Boreellus Aldrich & Shannon, 1923 5 subalpina (Ringdahl, 1931) atriceps (Zetterstedt, 1845) uralensis VilIeneuve, 1922 boganidae (Erichson, 1851) vicina Robineau-DesvoidY, 1830 aristatus Aldrich & Shannon, 1923 erythrocephala (Meigen, 1826), preocc. Protocalliphora Hough, 1899 I vomitoria (Linnaeus, 1758) A vihospita Hendel, 1901 Cynomya Robineau-Desvoidy, 1830 Orneocalliphora Peus, 1960 mortuorum (Linneaeus, 1758) azurea (Fallen, 1817) Lucilia Robineau-Desvoidy, 1830 sordida (Zetterstedt, 1838) Phaenicia Robineau-Desvoidy, 1863 chrysorrhoea (Meigen, 1826) Bufolucilia Townsend, 1919 nuortevai Grunin, 1972 Francilia Shannon, 1924 proxima Grunin, 1966 Acrophagella Ringdahl, 1942 Protophormia Townsend, 1908 5 bufonivora Moniez, 1876 terraenovae (Robineau-Desvoidy, 1830) caesar (Linnaeus, 1758) groenlandica (Zetterstedt, 1838) cadaverina (Linnaeus, 1758) Trypocalliphora Peus, 1960 splendida (Meigen, 1826) braueri (Hendel, 1901) fuscipalpis (Zetterstedt, 1845) hirudo (Shannon & Dobroscky, 1924) magnicornis (Siebke, 1863) lindneri Peus, 1960 alaskensis (Shannon, 1924) illustris (Meigen, 1826) simulatrix (Pandelle, 1896) regalis (Meigen, 1826) Subfamily Polleninae longilobata (Pandelle, 1896) richardsi Collin, 1926 Pollenia Robineau-Desvoidy, 1830 sericata (Meigen, 1826) angustigena Wainwright, 1940 6 silvarum (Meigen, 1826) griseotomentosa (Jacentkovsky, 1944) 7 Melinda Robineau·Desvoidy, 1830 varia: Wainwright 1932, Emden 1954, Xerophilophaga Enderlein, 1933 not Meigen (misidentification) cognata (Meigen, 1830) intermedia Macquart, 1835 caerulea (Meigen, 1826), preocc. nitens (Zetterstedt, 1845) 8 anthracina (Meigen 1838) vespillo: authors, not Fabricius (misi gentilis Robineau-Desvoidy, 1830 dentification) Onesia Robineau-Desvoidy, 1830 pseudorudis nom.nov. 9 Macrophallus Mueller, 1922 obscura Bigot, 1888, preocc., stat.n. Maeronesia Villeneuve, 1926 rudis (Fabricius, 1794) 10 sepulcralis (Meigen, 1826) obscura (Fabricius, 1794) 11 subapennina Rondani, 1862 vagabunda (Meigen, 1824)

90 Annotations spillo was erroneously assigned by Meigen (and subsequently by most modern authors, (I) Bellardia agilis (Meigen). Musca agilis Mei e.g. Zumpt 1956 and MihaIyi 1979) and for gen, 1826 is a junior primary homonym of which Zetterstedt introduced the name Mu Musca agilis Harris, 1780 (Tachinidae) sca nitens. The two specimens (probably (Crosskey 1975) syntypes) of Musca vespillo Fabricius, 1794 (2) Bellardia pusilla (Meigen). Musca pusilla present in Fabricius' collection (the Kiel col Meigen, 1826 is a junior primary homo lection) (cf. Zimsen 1964) in fact both be nym of Musca pusilla Gmelin, 1790 (Syrp long in Bellardia according to own observa hidae). tions, and the correctness of Zetterstedt's is (3) Pseudonesia (monotypic) should be inclu view thus confumed. Unfortunately Mu ded in Bellardia by virtue of phallosome sca nitens Zetterstedt, 1845 is a junior pri and ovipositor structure as well as external mary homonym of Musca nitens de Villers, features, according to own observations. 1789 (Syrphidae). (9) Among Pollenia material from Europe, (4) My reasons for treating Helicobosca as a North America and even New Zealand I calliphorid rather than a paramacronychi have seen numerous specimens belonging ine (agriine) sarcophagid as done by e.g. to an hitherto unrecognised species. It runs Verves (I980, 1982) and for assigning it to a to rudis in MihaIyi's key (I 976), but is easily separate subfamilY will be given in a fort distinguished from both rudis and angusti hcoming paper. gena (not considered by Mihalyi) in both (5) The monotypic genera Boreellus and Pro sexes by the presence of a bundle of pale tophormia should probably be merged. hairs on the underside of the wing at the Rognes (I985) cites a number of synapo junction of hand sc. The male can also be morphies common only to these genera, recognised by the absence on the middle and Collin (1925), Rodendorf (1926) and third of the hind tibia of any erect av hairs, S6guy (1928) regard the constituent species only normal decumbent setulae are present as congeneric. besides the av setae. In rudis males there are (6) Contrary to Wainwright (1940) and Pont numerous erect longish av hairs in addition (1975) I regard angustigena as a good spe to the av setae (usually less conspicuous in cies, not merely a subspecies of rudis. I do angustigena). Neither is the ventral vesti not accept the synonymisation made by ture of the male abdominal tergites in any Zumpt (I 955, 1956). way as dense and fine as in rudis males, but rather similar to the one present dorsally (as (7) British material of IIvariall agrees very well in angustigena). There are also constant and with Zumpt's (I956) and Mihalyi's (1976) distinct differences in the structure of the descripti0ns of griseotomentosa. The termi distiphallus from both rudis and angusti nalia of British and Norwegian material gena. I have recently been able to examine also agree with Mihalyi's (1979) figures. the d holotype of Pollenia obscura Bigot, Mihlilyi (1976) describes the basicosta as 1888: 597 from ((Amer. septentr.» (in black, but not all the specimens I have seen BMNH, London). It belongs to the present are that dark. This feature varies. Neither taxon. Bigot's name, treated as a synonym Wainwright (I932, 1940), Emden (I954) of rudis by Hall (1965), is a junior secon nor Mihalyi (1976, 1979) refer to the single dary homonym, preoccupied in Pollenia by ph seta (outer one lacking), an apparently Musca obscura Fabricius, 1794. I have used constant feature of this species. the proposed replacement name on determi (8) Pollenia nitens (Zetterstedt). Zetterstedt nation labels in various collections. (I 845: 1340) cites Meigen to the effect that (10) Nothing but an empty pin carrying a label Meigen's Musca sepuleralis (a species with reading ((rudis» in Fabricius' hand remains yellow palpi) according to Wiedemann is of the holotype of Musca rudis Fabricius, present in Fabricius' collection under the 1794: 314 in the Kiel collection in ZM Co name Musca vespillo. He also points out penhagen. A neotype corresponding to the that Fabricius' older name vespillo should concept of rudis of Wainwright (I 940) be used for this species and not for the Pol (very tight ventral abdominal vestiture of lenia species with black palpi and shining male, cf. above note 9) should be designa blackish abdomen to which the name ve- ted.

91 Cl 1) I have recently examined the d holotype of Emden, F.r. van 1954. Diptera Cyclorrhapha Ca Musca obscura Fabricius, 1794: 315 in the 1yptrata (I). Section (a) Tachinidae and Calliphori !Gel collection in ZM Copenhagen. It lacks dae. Handbk Ident. Br. Insects 10 (4) (a), 1...,...133. the head and all legs. It clearly belongs to Hackman, W. 1980. A check list of the Finnish Dip tera n. Cyclorrhapha. Notul. ent. 60,117-162. rudis in the sense of Wainwright (I 940). Hall, D.G. 1965. Family Calliphoridae. - Pp. 922-932, in: Stone, Sabrosky, Wirth, Foote & Coulson, A catalog oJthe Diptera oJAmerica north ACKNOWLEDGEMENTS oJMexico. Agric. Handbook 276: iv + 1696 pp., Many thanks are due to 1. Greve Jensen, Mu Washington D.C. Lundbeck, W. 1927. Diptera Danica. Part Vll. Platy seum of Zoology, Bergen, J.E. Raastad, Mu pezidae, Tachinidae. G.E.C. Gad, Copenhagen. seum of Zoology, Oslo, A. Fjellberg, TromS0 MihaIyi, F. 1976. Contribution to the knowledge of Museum, and J.O. Solem, DKNVS-Museet, the genus Pollenia R.-D. (Diptera: Calliphoridae). Trondheim, for loan of material; to S. Andersen, Acta zool. Acad. scient. Hung. 22, 327-333. Copenhagen, and AC. Pont, London, for gift - 1979. Femeslegyek - Huslegyek Calliphoridae and loan of Danish and British material, respec - Sarcophagidae. Magyarorszag allatvilaga tively, and for important information; to S. Funa Hungariae 15 (J 6), I -152 (Fauna Hunga, Andersen, 1. Lyneborg and V. Michelsen, Co riae 135, ISBN 963 05 2027 3). penhagen for a most useful stay at the Copenha Peus, F. 1960. Zur Kenntnis der ornithoparasitischen Phormiinen (Diptera, Calliphoridae). - Dt. ent. gen museum; to A.C. Pont for a most useful stay Z. (N.F.) 7, 193-235. at the British Museum (Natural History), Lon Pont, A.C. 1975.81. Calliphoridae. - In: Kloet, G.S. don; to B. Sagvolden and J.H. Simonsen for let & Hincks, W.D. (eds.l, A check list of British in ting me see their collections; and to B. Hliland, sects. Second edition. Part 5: Dipteta and Sipho T. Jonassen, F. Midtgaard, TR. Nielsen, A.-J. naptera. Handbk Ident. Br. Insects 9 (5), 107 Nilsen, J.H. Simonsen and S. Svendsen for gift -108. of precious material. The grant from Stavanger - 1981. The Linnaean species ofthe families Fannii- Lrererhegskole 1982-1983 is gratefully ack dae, Anthomyiidae and Muscidae (Insecta: Dipte owledged. A grant (D.68.40-023) from Norges ra). Bioi. J. Unn. Soc. 15,165-175. RingdahI, O. I 944a. Revision av Vilhe1m Storms Almenvitenskapelige Forskningsnid made pos Diptersamling r. Familjen Tachinidae. K. Norske sible the visits to Copenhagen and London. Vidensk. Selsk. Forh. 17 (I9J. 79-81. - 1944b. Contributions to the knowledge of the ta chinids and muscids of Norway. TromslJ Mus. Aarsh. 65 (I 942) (4), 1-27. REFERENCES - 1945. Forteckning over de av Zetterstedt i Insecta Lapponica och Diptera Scandinaviae beskrivna ta Ardo, P. 1957. Studies in the marine shore dune eco chiniderna med synonymer jiimte anteckningar system with special reference to the dipterous over en del arter. Opusc. ent. 10, 26-35. fauna. Opusc. ent. Suppl. 14, 255 pp. - 1952. Catalogus Insectorum Sueciae XI Diptera Aubertin, D. 1933. Revision ofthe genus Lucilia R. Cyclorrapha: Muscaria Schizometopa. Opusc. ent. D. (Diptera, Calliphoridae). J. Unn. Soc. Lond. 17, 129 - 186. Zool. 38, 389-436. Rodendorf, B.B. 1926. Morphologisches Studium an Bidenkap, O. 1892. UnderS0ge1ser over Diptera Bra ilusseren Genitalorganen der CalIiphorinen (Dip chycera i Jar1sberg og Laurvigs amt sommeren tera). Russkii Zoologicheskii Zhurnal (Revue zoo 1891. Ent. Tidskr. 13, 225-246. logique russe) 6 W, 83 -128. (In Russian, with - 190 I. Forel0big oversigt over de i det arktiske English title and summary). Norge hidtil bemrerkede Diptera Brachycera. Rognes, K. 1980. The blow-fly genus Lucilia Robi TromslJ Mus. Aarsh. 23 (J 900), 13 - 112. neau-Desvoidy (Diptera, Calliphoridae) in Nor Bigot, J.-M.-F. 1888. Dipteres nouveaux ou peu con way. Fauna norv. Ser. B. 27, 39-52. nus. Muscidi (J.B.). Bull. Soc. zool. Fr. 12 (5 -6) - 1982. A small collection of calypterate Diptera (I887), 581-617. (Published January 1888 accor (Tachinidae, Sarcophagidae, Calliphoridae, Mu ding to information from A.C. Pont, London). scidae) from the Dovre mountains, Southern Nor Collin, J.E. 1925. Diptera (Cyclorrhapha) from Spits way. Fauna norv. Ser. B. 29, 110-114. bergen. Results of the Oxford University Expedi - 1983a. Additions to the Norwegian fauna of ca tion to Spitsbergen 1924. Ann. Mag. nat. Hist. (9) 1ypterate Diptera (Tachinidae, Calliphoridae, Mu 16,332-337. scidae). Fauna norv. Ser. B. 30, 88-93. Crosskey, R.W. 1975.78. Tachinidae. - In: Kloet, - 1983b. The identity of Sarcophaga magnicornis G.S. & Hincks, W.D. (eds.), A check list of British Siebke, 1863 (Diptera: Calliphoridae). Ent. scand. insects. Second edition. Part 5: Diptera and Sipho 14, 405-406. naptera. Handbk Ident. Br. Insects 9 (5), 95 -105. - 1985. Revision ofthe bird-parasitic blowfly genus

92 Trypocal/iphora Peus, 1960 (Diptera: Calliphori dited and posthumously published by J. Sparre dae). Ent. scand. 15, 371-382. Schneider). - in press. On the identity of Sarcophaga alpina let Verves, Yu.G. 1980. Sostav podsemeistva Parama terstedt, 1838 and the interpretation ofAcrophaga cronychiinae (Diptera, Sarcophagidae) Starogo Brauer & Bergenstamm, 1891 (Diptera: Callipho Sveta. {The composition ofthe subfamily Parama ridae). Ent. scand. cronychiinae (Diptera, Sarcophagidae) of the Old Sch0yen, W.M. 1889. Supplement ill H. Siebke's World). Zool. Zh. 59,1476-1482. Enumeratio Insectorum Norvegicorum, Fasc. IV. - 1982. 64h. Sarcophaginae, pp. 235 - 296 (Liefe (Diptera). Christiania Vidensk. Selsk. Forh. 1889 rung 327). - In: Lindner, E. (ed.), Die Fliegen (12),1-15. der palearktischen Region, 11. Schumann, H. 1964. Revision der Gattung Onesia Wainwright, C.J. 1932. The British Tachinidae (Dip Robineau-Desvoidy, 1830. Beitr. Ent. 14, 915 tera). First supplement. Trans. ent. Soc. Lond. 80, -938. 405-424. - 1973a. Bemerkungen rum Status der Gattungen - 1940. The British Tachinidae (Diptera): Second Onesia, Melinda und Be/lardia (Diptera, Callipho supplement. Trans. r. ent. Soc. Lond. 90, 411 ridae). Mitt. zool. Mus. Berlin 49, 333-344. -448. - 1973b. Revision der palaearktischen Melinda-Ar letterstedt, J.W. 1845. Diptera Scandinaviae dispo ten (Diptera: Calliphoridae). Dt. ent. Z. (N.F.) 20, sita et descripta, 4, 1281 -1738. Lundae l 293-314. ( = Lund). - 1974. Revision der palaearktischen Be/lardia-Ar limsen, E. 1964. The type material ofre. Fabricius. ten (Diptera, Calliphoridae). Dt. ent. Z. (NF) 21, Munksgaard, Copenhagen. 231-299. lumpt, F. 1955. The width offrons as a variable fea Seguy, E. 1928. Etudes sur les mouches parasites. ture in the Calliphorinae (Diptera), with descrip Tome I Conopides, Oestrides et Calliphorines de tions oftwo new species. J. ent. Soc. S. Africa 18, l'Europe occidentale. Recherches sur la morpho 50-59. logie et la distribution geographique des Dipreres - 1956. 64i. Calliphorinae. - In: Lindner, E. (ed.), it larves parasites. Encycl. ent. (A) 9, 1-251. Die Fliegen der palaearktischen Region 11, Siebke, H. 1877. Enumeratio Insectorum Norvegico 1-140 (Lieferungen 190, 191, 193). rum. Fasciculum IV. Catalogum Dipterorum Con tinentem. xiv + 255 pp. Christianiae (= Oslo). (E- Received 17 Aug. 1984

f I

93 .. 1 A check-list of Norwegian Fanniidae (Diptera)

KNUT ROGNES

Rognes. K. 1985. A check-list of Norwegian Fanniidae (Diptera). Fauna norv. Ser. B. 32. 94-96.

Fanniid material from Norway in the museum collections of the Universities in Bergen, Oslo and Troms0 and in my own collection has been revised and a check-list of 42 species of Fannia Robineau-Desvoidy known at present from Norway is presented. Fannia bigelowi ChiIlcott, 1961 is recorded from the Palaearctic Region for the first time. Knut Rognes, Stavanger Lererh0gskole. Postboks 2521 Ullandhaug. N-4001 Stavanger. Norway.

The check-list of Fanniidae presented below is 1975, 1983). Undoubtedly many species still based upon material from Norway in the mu· await discovery in Norway. seum collections of the Universities in Bergen The nomenclature used follows the one of (mainly O. Bidenkap, A. Fjellberg, L. Greve, A. A.C. Pont in his Fanniidae contribution for the L0ken, N. Knaben, T.R. Nielsen leg.), Oslo (ma forthcoming Catalogue of Palaearetic Diptera (e inly J. Knaben, R. Krogerus and J.H.S. Siebke ditor A. S60s, Budapest). Specific synonyms leg.) and Troms0 (mainly O. Ringdahl, T. Soot have been included when based on type-speci Ryen leg.) about 700 specimens) and in my mens from Scandinavian localities, or when ne own collection (about 800 specimens). In addi cessary for species recognition. In the list all sy tion a few specimens in the private collections of nonyms have been indented relative to valid K. Berggren, B. Sagvolden and J.H. Simonsen names. Superscript numbers refer to annota have been examined. I have not seen the V. tions at the end of the list. Storm collection in Trondheim revised by Ring dahl (l944a). Some unidentified material collec ted by P. Ardo, H. Andersson and R. Dahl in Family Fanniidae Northern Norway has also been examined (ma terial in Lund). I have examined specimens from Fannia Robineau-Desvoidy, 1830 Norway of all the species listed except 2. My Homalomyia Bouche, 1834 reasons for listing these are given in the annota Coelomyia Haliday in Westwood, 1840 tions at the end of the list. Ivalomyia Tiensuu, 1938 Ringdahl (1952) listed 21 species of Fanniidae armata (Meigen. 1826) from Norway, but did not include Fannia nor· atra (Stein, 1895) vegica described from Troms0 (Ringdahl 1934). bigelowi Chillcott, 1961 1 His list summarised previous work into the Nor canicularis (Linnaeus, 1761) wegian fauna (Ringdahl 1928, I 944a, 1944b) lateralis (Linnaeus, 1758) (suppressed) and unpublished examination of parts of the carbonaria (Meigen, 1826) Oslo collection. Ringdahl (1954) added I species carbonella (Stein, 1895) and Rognes (1982) 1, bringing the total number coracina (Loew. 1873) known from Norway to 24. The present revi corvina (Verrall. 1892) sion lists 42 species from Norway. In compari halterata Ringdahl, 1918 son it may be noted that 39 species have been cothurnata (Loew, 1873) 2 published from Finland (Hackman 1980), 50 difficilis (Stein, 1895) species from Sweden (Ringdahl 1952, Rognes !uscu{a (FalUm, 1825) 1982) and 60 species from the BritiSh Isles (Pont genualis (Stein, 1895) glaucescens (Zetterstedt, 1845) hirticeps (Stein, 1892) 1) Paper no. 3 in a series based on investigations fi incisurata (Zetterstedt, 1838) nanced by a grant (<

94 FaufUJ no"'. Ser. B. 32: 94-96. Oslo 1985. lugens (Zetterstedt, 1845) I have compared the specimens with Nearc mutica (Zetterstedt, 1845) tic material (including 2 d d paratypes) in limbata (Tiensuu, 1938) the CNC, Ottawa. The cercal plate which is lugubrina (Zetterstedt, 1838) illustrated by Chillcott (961) is very cha albinervis (Zetterstedt, 1845) racteristic. The Norwegian specimens and lustrator (Harris, 1780) the paratypes examined lack postoccipital hamata (Macquart, 1835) setulae in the upper part of the occiput. viduata (Zetterstedt, 1845) (2) Fannia cothurnata Loew, 1873. Not seen. zetterstedti (Bidenkap, 1892) Included on the basis of record in Ringdahl manicata (Meigen, 1826) 0928:26, «Von Strand bei Ledingen gefun armillata (Zetterstedt, 1838) den. Bisher nicht mit sicherheit anderswo in melania (Dufour, 1839) Skandinavien gefunden.»). Ringdahl may ciliata (Stein, 1895) not have seen Strand's specimen(s), but minutipalpis (Stein, 1895) Strand 0906: 104) reports the identification mollissima (Haliday in Westwood, 1840) to have been made by P. Stein. spathulata (Zetterstedt, 1845) (3) Fannia norvegica Ringdahl, 1934. Not faroensis (Lyneborg, 1962) seen. Holotype d taken at Troms0, Nort monilis (Haliday, 1838) hern Norway, 10 July 1926 (Ringdahl norvegica Ringdahl, 1934 3 1934). pallitibia (Rondani, 1866) parva (Stein, 1895) polychaeta (Stein, 1895) postica (Stein, 1895) parapostica Hennig, 1955 ACKNOWLEDGEMENTS posticaria d'Assis Fonseca, 1968 (una I would like to thank L. Greve Jensen, Museum vailable) of Zoology, Bergen, J.E. Raastad, Museum of posticata (Meigen, 1826) Zoology, Oslo and A. Fjellberg, Troms0 Mu pretiosa (Schiner, 1862) seum, for having made the collections of calyp rondanii (Strobl, 1893) terate Diptera in their care available to me; A.c. aerea: authors, not Fallen (misidentifi Pont, London, for most generously having put cation) his ms copy ofhis Fanniidae contribution for the scalaris (Fabricius, 1794) Catalogue of Palaearctic Diptera at my disposal serena (Fallen, 1825) and for having checked some of my identifica similis (Stein, 1895) tions; J.R. Vockeroth, Ottawa, for the loan of sociella (Zetterstedt, 1845) material in the CNC; K. Berggren, B. Sagvolden spathiophora Malloch, 1918 and J.H. Simonsen for letting me see their collec no;iulosa Ringdahl, 1926 tions; and to B. Borgersen, T. Jonassen, F. Midt stigi Rognes, 1982 gaard, T.R. Nielsen, A.-J. Nilsen and S. Svend subatripes d'Assis-Fonseca, 1967 sen for gift of specimens. The grant from Sta ,subpellucens (Zetterstedt, 1845) vanger l.

REFERENCES Chillcott, J.G. 1961. A revision of the Nearctic spe Annotations cies of Fanniinae (Diptera: Muscidae). Can. Ent. (I) Fannia bigelowi Chillcott, 1961. There are Supp/. 14 (J960), 1-295. two specimens in the Oslo collection: Opp Hackman, W. 1980. A check list of the Finnish Dip land, ON, Sel: Uirgard 1 d 27 June 1861 tera. 11. Cyc1orrhapha. Notu/. Ent. 60, 117 -162. Pont, A.C. 1975. 84. Fanniidae. - In: Kloet, G.S. & (Siebke) (ZMO no. 8395) and ON, Dovre: Hincks, W.D. (eds.), A check list of British In Fokstua 1 d 17 July 1953 (Krogerus) (ZMO sects. Second edition. Part 5: Diptera and Sipho no. 10272), both EIS 71. These are the first naptera. Handbk Ident. Br. Insects 9 (5), 113 ones recorded from the Palaearctic Region. -114.

95 - 1983. Fannia lineata (Stein, 1895), new to Britain Aarsh. 65 (]942), (4), 1-27. (Diptera, Fanniidae). Entomologist's mono Mag. - 1952. Catalogus insectorum Sueciae XI Diptp.ra 1l9,229-231. Cyclorrapha: Muscaria Schizometopa. Opusc. ent. Ringdahl, O. 1928. Beitrage zur Kenntnis der Antho /7,129-186. myidenfauna des nordlichen Norwegens. Tromse - 1954. Nya fyndorter for norska Diptera. Norsk Mus. Aarsh. 49 (926), (J), 1-60. ent. Tidsskr. 9, 46 - 54. - 1934. Einige neue Musciden. Konowia 13, 97 Rognes, K. 1982. Fannia stigi n.sp. from Scandinavia -100. (Diptera: Fanniidae). Ent. scand. 13, 325-330. - 1944a. Revision av Vilhelm Storms Diptersamling Strand, E. 1906. Nye bidrag ill Norges hymenopter 2. Familjen Muscidae. K. norske Vidensk. Selsk. og dipterfauna. Ill. Nye norske 10kaliteter for Forh. 17 (20). 82-85. Diptera. Nyt. Mag. Naturvid. 44, 102-104. - 1944b. Contributions to the knowledge of the ta chinids and muscids of Norway. Tromse Mus. Received 17 Aug. 1984

96 I ~ Additions to the Norwegian fauna of Dolichopodidae (Dipt.)

TERJE JONASSEN

Jonassen, T. 1985. Additions to the Norwegian fauna of Dolichopodidae (Dipt.). Fauna norv. Ser. B, 32, 97-99.

Fourteen species of Dolichopodidae are reported from Norway probably for the first time.

Terje Jonassen, N-4170 Sjernar0Y, Norway.

INTRODUCTION When Siebke published his Enumeratio Insecto Dolichopus atripes Meigen, 1824 rum Norvegicorum, Fasc. IV in 1877, he listed RI, Forsand: R0ssdalen, EIS 7, I c5 26 June 108 species ofthe family Dolichopodidae as part 1982; Songesand, EIS 7, IQ 18 July 1984; of the Norwegian fauna. Discounting the ob Daladalen, EIS 8, I c5 26-27 July 1984. vious errors of Siebke's list, e.g. his description Dolichopus caligatus Wahlberg, 1850 ofconspecific males and females under different RI, Forsand: Helmikst01, EIS 8, I c5 5 August names, the total number will be reduced to 1982. An uncommon species with a sparse about one hundred. Nevertheless, Siebke's col distribution in northern Europe. This speci lection still make up the great bulk ofthe known men was captured among marsh vegetation. Norwegian material of Dolichopodidae, new re later searches for this particular species at cords have been few and scattered. However, the same location have proved fruitless. species new to the Norwegian fauna were added Dolichopus wahlbergi Zetterstedt, 1843 by Sch0yen (1889) - I sp., Bidenkap (1900 RY, Sandnes: Gravaren, EIS 7, 2 c5 c5, I Q I sp., Frey (1914) - 2 spp., Ringdahl (1928, 30 July 1982. The specimens were taken 1954) - 8 spp., Thuneberg (1955) - I sp. and among shady foliage in a small decidious Hedstrom (I975) - I sp. wood growing on clayey soil. Below, ·14 species of Dolichopodidae are re Hercostomus cupreus (Fallen, 1823) ported from Norway probably for the first time. RY, Sandnes: H01e, EIS 7, I c5, 2 Q Q 20 The material for this report has been collected June 1981, I c5 4 June 1982; Ims, EIS 7, IQ by the author and is deposited in the author's 4 June 1982; Dale, EIS 7, 2 c5 c5 9 June 1982; collection, when nothing else is mentioned. The Melshei, EIS 7, IQ 14 June 1982; Klepp: methods ofcollecting have been the same as spe 0ksnevad, EIS 7, I Q 15 June 1982; Sta cified in Jonassen (I 984). vanger: Madlaforen, EIS 7, I c5 4 July 1981, The arrangement ofsubfamilies and sequence K. Rognes; Rennes0Y: Vikevag, EIS 14, I c5 ofspecies largely follow C.E. Dyte's list in Kloet 24 May 1981, 2 c5 c5, IQ 30 May 1981, and Hincks (1975). 6 c5 c5, 4 Q Q 10 June 1982; Morkavatn, EIS The geographical division of the districts fol 14, IQ II June 1982; F0rsvoll, EIS 14, I c5 lows 0kland (I981). II June 1982; RI, Forsand: Songesand, EIS 7, I c5, IQ 14 June 1981, IQ 13 June 1981, I c5 19 June 1983, I c5 29 June 1983; R0SS dalen, EIS 7, 2 c5 c5 26 June 1982. Males are SYSTEMATIC LIST easily recognized because of the particular thickening and chaetotaxy of the mid tibia. Subfamily Dolichopodinae This is one ofthe most common Hercostomus Dolichopus acuticornis Wiedemann, 1817 species in Rogaland. RY, Ha: Brusand, EIS 3, 4 c5 c5, 7 Q Q 7 Hercostomus nigrilamellatus (Macquart, 1827) July 1982. A typical sea coast species. These VAY, Vennesla: Frikstad, EIS 5, I c5 21 June specimens were captured among low her 1983, B. Borgersen. This specimen was bage between dunes on a dry, sandy beach. found in a hollow ash (Fraxinus excelsior L.).

Fauna norv. Ser. D, 32, 97-99. Oslo 1985. 97 .. According to the collector, Mr. Bj0rnar Bor 7, 1 Q 20 June 1981, K. Rognes; Klepp: 0ks gersen (in litt.), the fly was probably reared in nevad, EIS 7,1 d 15 June 1982; RI, Forsand: this said hollow, since it appeared among the Songesand, EIS 7, I d 19 June 1983, 1 d 20 sieved detritus from the cavity of the tree, June 1983. An examination ofthe specimens and because the specimen was immature. under the name Hydrophorus (= Medetera) Hypophyllus obscurellus (Fallen, 1823) muralis Meigen, 1824 in the still existing RY, Rennes0Y: Vikevag, EIS 14, Id 6 Sep parts of Siebke's collection in ZMO, will pos tember 1981, 5 dd, 3 Q Q 25 July 1982, sible yield some infumata. The genitalia of 1 d, 1 Q 26 August 1983. All specimens the four males in my collection have been were captured on a wet, vertical rock close to dissected and agree with the figures given by a water-fall. Negrobov & Stackelberg (1974).

Subfamily Hydrophorinae Subfamily Neurigoninae Hydrophorus albiceps Frey, 1915 Neurigona quadrifasciata (Fabricius, 1781) RY, Rennes0Y: Sel, EIS 14, IQ 18 April OS, S0r-Fron: Harpefoss, EIS 62, I Q 16 July 1982; RI, Forsand: Songesand, EIS 7, 1 d 22 1982, 0. & K. Rognes. August 1981, 3 Q Q 26 March 1983; Songe sandst0len, EIS 7, 2 d d 20 August 1982; Helmikst01, EIS 8, 1 d, I Q 5 August 1982, Subfamily Campsicneminae 3 dd, I Q 5 September 1982. Even though Campsicnemus (Ectomus) alpinus (Haliday, this species has not been mentioned in pre 1833) J vious Norwegian records, it is highly pro RI, Forsand: Songesandst0len, EIS 7, 1 Q 20 bably that an examination of Siebke's collec August 1982, Id 30-31 July 1984. tion in ZMO will yield some specimens of Chrysotimus concinnus (Zetterstedt, 1843) this species among those determined by him RY, Rennes0Y: Vikevag, EIS 14, 2 dd, 1 Q as Hydrophorus bipunctatus Lehmann or H. 13 July 1982, 1 d 14 July 1982, 1 Q 15 July binotatus Fallen. 1982. The species occurred in numbers among sun-exposed decidious trees and lo wer herbage during the short period which Subfamily Mediterinae the above dates indicate. A rather uncom Medetera excellens Frey, 1909 mon species with a sparse Scandinavian dis RI, Forsand: Songesand, EIS 7, 2 d d 14 tribution. August 1982. A rather rare species, described from Finnish specimens. Elsewhere it seems to be known only from 210calities in Sweden (Ringdahl 1950, 3 Scottish specimens (Fon ACKNOWLEDGEMENTS seca 1978), from East Germany and from the Leningrad area in the Soviet Union (Negro My sincere thanks go to Mr. Knut Rognes, bov & Stackelberg (1972». Madla, and to Mr. Bj0rnar Borgersen, 0stre Hal Medetera fasciata Frey, 1915 sen, for the gifts of precious material. RI, Forsand: Songesand, EIS 7, 1 d 16 May 1983; 1 d 4 June 1983, 2 dd, 1 d Q (in co pula) 19 June 1983. These are the western most finds ofthis species from the Palaearctic REFERENCES region. Previously known from Sweden, Fin Bidenkap, O. 190 I. Fore10big oversigt over de i det land and the USSR (Negrobov & Stackelberg arktiske Norge hidtil bemrerkede Diptera Brachy 1972). The genitalia have been dissected in cera. TromS0 Mus. Aarsh. 23 (1900),13-112. four ofthe male specimens, and they all agree Fonseca, E.C.M.d'Assis. 1978. Diptera: Dolichopodi dae. Handbk. Ident. Br. Insects IX (6), 1-90. with the figures given by Negrobov & Stack Frey, R. 1914. Verzeichnis von Dr. E. Strand in Nor elberg (1974). wegen gesarnmelter Diptera Brachycera (Faro. Medetera infumata Loew, 1857 Empididae, Dolichopodidae und Lonchopteridae). RY, Sandnes: Ims, EIS 7, 1 Q 4 June 1982; In: Strand, E. Neue Beitriige zur Arthropodenfa Melshei, EIS 7, Id, I Q 14 June 1982, IQ 6 una Norwegens, part XVII. Nyt Mag. Naturvid. July 1982; Strand: towards Prekestolen, EIS 51,320-322.

98 Hedstrom, L. 1975. Dolichopodidae (Diptera). Fauna Ringdahl, a. 1954. Nya fyndorter fOr norska Dip of the Hardangervidda, No. 9 (1976), 13 - 21. tera. Norsk ent. tidsskr. 9 (1953),46-54. Jonassen, T. 1984. Some recent records of Hybotidae Sch0yen, W.M. 1889. Supplement til H. Siebke's and Microphoridae (Dipt., Empidoidea) from Enumeratio Insectorum Norvegicorum, Fascicu Norway. Fauna norv. Ser. B. 31,92-95. lum IV. (Diptera). Christiania Vidensk. Selsk. Kloet, G.S. & W.D. Hincks (eds.). 1975. A check list Forh. 1889 () 2), 1-15. of British insects. Second edition. Part 5: Diptera Siebke, H. 1877. Enumeratio lnsectorum Norvegico and Siphonaptera. Handbk. Ident. Br. Insects XI rum. Fasciculum IV. Catalogum Dipterorum (5). Continentem. A.W. Br0gger, Christiania, 255 pp. Negrobov, a.p. & A.A. Stackelberg. 1972. 29. Doli Thuneberg, E. 1955. A revision of the Palaearctic chopodidae pp. 238-256. In: E. Lindner (Ed.); species of the genus Medetera Fischer (Dipt., Do Die Fliegen der palaearktischen Region 4 (5). lichopodidae). Suom. hyont. Aikak. 21, Negrobov, a.p. & A.A. Stackelberg. 1974. 29. Doli 130-157. chopodidae pp. 257 - 302. In: E. Lindner (Ed.): 0kland, K.A. 1981. Inndeling av Norge til bruk ved Die Fliegen der palaearktischen Region 4 (5). biogeografiske oppgaver - et revidert Strandsys Ringdahl, a. 1928. Neue skandinavische Dipteren. tern. Fauna, Oslo 34,167-178. Ent. Tidskr. 49, 18 - 21. Ringdahl, a. 1951. Flugor fnin Lapplands, Jiimtlands Received 8 Jan. 1985. och Harjedalens fjiilltrakter (Diptera Brachycera). Opusc. Ent. 16, 113-186.

; f

99 Distribution and food habits of mayflies (Ephemeroptera) in streams in the Dovrefjell mountains, Central Norway

TERJE NOST

Nbst, T. 1985. Distribution and food habits of mayflies (Ephemeroptera) in streams in the Dovrefjell mountains, Central Norway. Fauna narY. Ser. B. 32. 100-105. Mayflies dominated the lotic communities in the subalpine and low alpine zones, although there was a marked decrease in number of individuals and species with increasing altitude. Baetis rhadani was the dominant species. Mayflies were absent above the low alpine zone. All species were herbivorous, feeding on ultrafine particulate organic matter (UPOM) in the subalpine and shifting to a mixture ofalgae and UPOM in the low alpine. This change in the diet is discussed in relation to available food items and their nutritive value. The results indi cates that temperature rather than food resources is the major factor determining distribu tion of mayflies in the area. Terje Nost, University ofTrondheim, The Museum, Erling Skakkesgt. 47A, N-7000 Trond- heim, Norway. J

INTRODUCTION STUDY AREA During the last two decades a number ofstudies The Dovrefjell mountains (Fig. 1) in Central have been made on Norwegian Ephemeroptera Norway cover about 1 100 km2 with altitudes from lotic habitats. The studies have focused ranging from about 500 m up to 2 200 m a.s.l. mainly on distribution and relatively few have The geology is rather complex because two large included ecological aspects, although papers by regions within the southern Scandinavian Cale Elliott (1965), Larsen (1968), Steine (1972), Brit donids meet in this area. The border approxima tain (1975), Lillehammer & Brittain (1978), tely follows the river Driva. The western side of Baekken0980, Raddum & Fjellheim (1982) the valley have Precambrium high-grade gneis should be mentioned. The aim ofthis paper was ses and schists, the eastern side Cambro-silurian to compare the distribution of mayfly species medium-grade mica schists and greenstones. and their feeding habits in the subalpine, low al The climate of the area is rather continental, pine and mid-alpine zones in the Dovrefjell mo the central parts having an annual precipitation untains. Previously, only Lillehammer & Brit of 473 mm (Kongsvoll 900 m a.s.u. The annual tain (1978) have studied the vertical zonation of mean temperature is -O.loC (Hjerkinn 959 m mayflies in Norway. Baekken (I981) examined a.s.L) and only 19 days a year have daily mean food habits of Baetis rhodani Pictet, but the spe temperature above 10°C (Nordhagen 1943). The cimens were all sampled from the subalpine. subalpine zone reaches an elevation of Several authors (e.g. Gledhill 1960, Egglis 1 000-1 100 m a.s.l. The border between low haw 1969, Minshall & Kuehne 1969, Woodall & alpine and mid-alpine zones is about Wallace 1972, Allan 1975, Stoneburner 1977, 1 300-1 450 m a.s.l. (Gjrerevoll 1975). Rabeni & Minshall 1977, Lillehammer & Brit The investigation was carried out in the river tain 1978, Macan 1980, Raddum & Fjellheim Driva and the watershed ofits large tributarities, 1982) have investigated distribution of mayflies the streams Amotselva, Grovu and Vinstra. in lotic habitats. Factors considered important Driva, Amotselva and Grovu originate in the for their available food distribution are: latitude, mid-alpine zone, while Vinstra originates in the altitude, allochthonous material, water chemi low alpine zone. The streams are relatively fast stry, temperature and substrate. flowing, but the upper parts of Amotselva and a Mayflies are largely herbivorous, although a few sampling sites in the subalpine zone in few are omnivorous (Brittain 1982). Grovu, have stretches more lentic in character.

100 Fauna no",. Ser. B. 32: 100-105. Oslo 1985. Driva

oL' ----le10 km

Fig. 1. The Dovrefjell mountain area, the main rivers and tributarities (watershed marked).

A few samples were also taken at outlets oflakes The food habits ofthe nymphs were determi in the Grovu system. ned by stomach analysis based on techniques Allochthonous organic matter appeared to be described in Shapas & Hilsenhoff Cl 976). Fore much more abundant in the subalpine than the guts of nymphs were removed and the contents low and miq,.alpine zones. No obvious decrease were squeezed on to a glass slide and cleared in in amounts of algae towards higher altitudes lactic acid. Five randomly chosen transects was apparent. across the central portion of each mount were The .~ange in chemical parameters was: pH measured. Along the transects every food par 6.4-7.3, Ca++ 0.5-13.5 mg 1-1, KI8 ticle was counted and given an approximate per 5-87 pm hos (N6st 1981). Water temperatures cent by volume. The food particles were grou rarely exceeded 10°C in the lower parts and ped into two main categories; I. detritus and 2. 3- 5°C in the highest located sites. algae. The classifIcation of detritus was defIned into fIne particulate organic matter (FPOM 50 pm -I mm) and ultrafIne particulate organic matter (UPOM 0.5-50 pm) (Cummins & Klug MATERIAL AND METHODS 1979, Vannote et al. 1980). Algae were separa Nymphs ofmayflies were collected mainly from ted into diatoms and other algae. Food analysis

I June to August 1979 and 1980, but a few samp was restricted to nymphs with wing pads mode I· les were also taken in July 1984. A total of rately to nearly fully developed because of their 17 548 nymphs were sampled using the kick dominance in the samples and their size. method (Frost et al. 1971). The mesh size of the The closely related species Baetis fusca net used was 0.5 mm. At 54 sites 89 samples tus/Baetis scambus and Baetis vernus/Baetis each covering 5 min. of sampling time were ta subalpinus are difficult to separate in nymphal ken. The samples were fIxed in 70 % ethanol. stage, and they were treated as «single» species.

101 Table 1. Occurrence and relative abundance (%) of ephemeropteran species in the subalpine and low alpine zones in the main watercourses

Driva Amotselva Gr0VU Vinstra subalpine low alpine subalpine low alpine subalpine low alpine subalpine low alpine Ameletus inopinatus 0.1 1.7 13.3 0.2 0.4 5.6 Parameletus chelifer 0.7 Siphlonurus lacustris 0.8 30.1 12.4 Baetis lapponicus 0.9 0.2 25.8 14.9 0.7 23.0 29.7 Baetis macani 0.1 1.2 31.3 Baetis rhodani 98.3 99.8 65.4 34.3 44.8 18.8 75.9 64.3 BaetisJuscatus Iscambus 0.1 3.0 Baetis vernus Isubalpinus 0.3 6.6 34.3 15.4 37.5 Baetis spp. 0.7 2.3 Heptagenia dalecarlica <0.1 1.9 Leptophlebia vespertina <0.1 0.1 Ephemerel/a aurivillii 0.3 0.3 0.8 0.7 0.6 Number of individuals 11912 1318 1215 756 1356 16 544 431 Number of samples 20 9 10 10 18 4 6 2

RESULTS B. macani in Grovu, made up more than 50% Distribution of the mayfly nymphs. In each of the remaining six genera only one species was recorded. Two Mayflies dominated the invertebrate fauna at all of these (Ephemerella aurivillii and Ameletus in sites in the subalpine and low alpine zones. In opinatus) occurred at several sampling sites, the the river Driva and the stream Grovu, there was other at only a few. Except for Siphlonurus lacu a marked decrease in the importance ofmayflies stris which was collected in large numbers at when going from the subalpine to the low alpine one sampling site in the subalpine in Grovu, all zone. The opposite trend was found in the the remaining species had low densities. stream Vinstra, as more suitable substratum for The highest number of specimens was recor mayfly nymphs occurred at 1 000 m or higher ded in the river Driva, while highest number of elevations (Nost 1981). The upper limit for the species was recorded in the stream Grovu. distribution of mayflies in this area is about 1 350 m a.s.l., which coincides with the border between low and mid-alpine zones. Other aquatic invertebrates of importance in Food habits the streams were stonefly (Plecoptera) nymphs A total of 527 nymphs were analyzed for their and Chironomidae larvae. The latter accounted food habits, all rivers at different elevation being for 80-90% of numbers in the lotic commu represented. nity of the mid-alpine. In the subalpine zone the different mayfly Seven genera and a minimum of 11 species of species had very similar food habits (Tab. 2). All mayflies were recorded (Tab. 1). All species oc nymphs fed on detritus, primarily ultrafine par curred in the subalpine, but only seven represen ticulate organic matter (UPOM). With a few ex ting four genera were present in the low alpine. ceptions UPOM accounted for 90% or more in Baetis was the dominant genus with at least five volume ofthe ingested material. Fine particulate species in the subalpine and at least four in the organic matter (FPOM) was eaten by 50% or low alpine. B. rhodani was the most abundant more ofthe nymphs, except B. rhodani in which species sampled in Driva and Vinstra, and acco the equivalent value was only 30 %. FPOM ra unted for almost 100 % oftotal mayfly numbers rely made up more than lO % of the volume of in Driva and 65-75% in Vinstra. In Amot the gut contents. Algae were consumed in small selva and Grovu the dominance of B. rhodani amounts by a few nymphs ofS. lacustris. B. lap was not so apparent and a marked decrease in ponicus and B. vern us Isubalpinus. Diatoms and its importance with altitude was found. In the fJ1amentous algae occurred in all species except low alpine B. vernuslsubalpinus and B. lapponi S. lacustris in which only fJ1amentous algae cus in Amotselva and B. vernuslsubalpinus and were found.

102 In the low alpine zone algae were a conspicu ous and important part of the diet of all species (Tab. 2). The nymphs of all species showed a +1 +1 +1 +1 +1 0'<1" Or- great diversity in their feeding habits. Some ap 66r-:~ t"'j peared to specialize on algae, others on UPOM. -N-N N However, a mixture of detritus and algae was most common. There were no significant diffe rences in food preferences between sampling tl +1 +1 +1 +1 t-O\O"'l:j- ""=::t sites or different watercourses. Algae represen l..C>oOMr-: \0 <""'"IN N ted on average about 50 % ofthe ingested mate rial in B. rhodani and B. vernus!subalpinus, and ~ o : about 30 % in B. lapponicus. Diatoms and other it +1 +1 +1 +1 +1 +1 algae were of equal importance in the first two o r-:~~~ r species, while other algae dominated in B. lap o \C~N'o:::f'" 0\ ponicus.

N-ON 0'\ O..oM~ 0' N--- +1 +1 +1 +1 +1 r-:'""':~~ r \0\000- 0 \C\Ov-.::::t- """ DISCUSSION Mayflies dominated the invertebrate fauna of10 r-- 00 ~ tic communities both in the subalpine and low N- 0 alpine zones. Mayflies were absent above the +1 +1 tl r-- ~ low alpine zone. There are however records of 00 0 mayflies in the mid-alpine zone in southern Nor way, both in 1entic and lotic habitats (Brittain 1974, Lillehammer & Brittain 1978, Nost 1983, +1 +1 Lillehammer 1984). <""'"l <""'"l The ephemeropteran fauna of the subalpine o 0 and low alpine zones of the Dovrefjell mounta ins was dominated by B. rhodani which is a :E :;:6~;::i~~~ 6 o N common feature of mountain streams in nort it +1 tl +1 tl +1 +1 +1 +1 +1 +1 hern Scandinavia (U1fstrand 1975, Koksvik NO\Co\OMOOOOOO 6~r-:--CV)N--CV)N6~ 1979, Lillehammer & Brittain 1978, Nost 1983). -N There is a greater uniformity of substratum r-r-Nr--NONtn 00 and discharge in the Driva and Vinstra than in ~oO";o'r--:~o\oO~ o N the other watercourses. This partly explains the +! +1 +1 +1 +1 +1 +1 +1 +1 +1 differences in species composition between the OOOt-MOr-Nr-OOO ~M""':Nv)r---:M~oOO...o streams. Species, such as B. macani. S. lacustris ooO\O\O\ooO\O\O'.O'\O\r- and Leptophlebia vespertina were found in more slow-flowing parts or in lake outlets in the "8 streams Amotselva and Grovu. ..2 Studies on Ephemeroptera and Plecoptera at 'o different altitudes in Ovre Heimdalen, about 100 .gt:: km SW of Dovrefjell (Brittain 1974, Lilleham ''=0 mer & Brittain 1978, Lillehammer 1984), indi ~oo cate that temperature and food are the main fac 0. E +1 tors determining species distribution. In Eng o Vl Uta land, Minshall & Kuehne (1969) reported that ';:1 N"O water chemistry, especially low pH and bicarbo V·- -;> oD . nate ion, could be a limiting factor for Epheme 0:1"0 f- .S roptera at higher altitudes. Observations made by Stoneburner (1977) in the United States indi cate similar trends as in England. In rivers in

103 western Norway Raddum & Fjellheim (I982) ACKNOWLEDGEMENTS showed B. rhodani to be absent at pH below 6.0. I am greatly indebted to J.O. Solem and J.E. However, in the Dovrefjell mountains pH is al Brittain for valuable comments and linguistic ways above this level (N6st 1981) and pH is not corrections of the manuscript. The stUdy was therefore a limiting factor. The distribution of supported by the Ministry of Environment. mayflies should therefore be considered in rela tion to food resources and temperature. In the subalpine zone birch trees and dwarf willow (Salix) vegetation give rise to a conside rable input of terrestrial organic matter to the REFERENCES streams. Here mayflies feed on detritus. Above Allan, J.D. 1975. Faunal replacement and longitudi the subalpine zone the allochthonous input is nal zonation in an alpine stream. Verh. Int. Ver. considerably reduced. This is especially obvious Theor. Angew. Limnol. /9,1646-1652. in the mid-alpine zone, in which no continuous Baekken, 1. 1981. Growth patterns and food habits plant cover exists. The high percentage of algae of Baetis rhodani, Capnia pygmaea and Diura nanseni in a West Norwegian river. Holarct. Ecol. in the diet of mayflies in the low alpine zone is 4.139-144. probably a result of the reduced input of terre Brittain, J.E. 1974. Studies on lentic Ephemeroptera strial organic matter into streams, but may also and Plecoptera of Southern Norway. Norsk ent. indicate a need for food of high quality. Within Tidsskr. 2/,135-154. the range of food available, there is evidence of - 1975. The life cycle of Baetis macani Kimrnins selection, as growth of mayflies may be influen (Ephemerida) in a Norwegian mountain biotope. ced by different foods (McCullough et al. 1979, Ent. scand. 6, 240-244. Cianciara 1980). The high nutritive values of li - 1976. Experimental studies on growth if Leptoph (Ephemerop~ra). ving algae was demonstrated by Cummins & lebia vespertina (L.) Freshwat. Bioi. 6. 445-449. (I Klug 979), and they reported differences ofse - 1982. Biology ofmayflies. Ann. Rev. Entomol. 27, veral hundred Calories per gram between dia 119-147. toms and detritus. Utilized energy bound up in Brown, D.S. 1960. The ingestion and digestion of al algae may, however, differ, depending on spe gae by Cloeon dipterum L. (Ephemeroptera). Hy cies of algae and mayfly nymph (Brown 1960, drobiol. /6, 81 - 96. 1961, Cianciara 1980). The fact that there was - 1961. The food of larvae of Cloeon dipterum L. an abrupt shift in the food habits leads to the and Baetis rhodani (Pietet) (Insecta: Ephemeropte conclusion that algae are of high nutritive value ra). J. Anim. Ecol. 30, 55-75. for mayfly species in the area. However, despite Cianciara, S. 1980. Biometric and bioenergetic cha racterization of the development of Cloeon dipte there being no obvious reduction of algae in the rum (L.). Pol. Arch. Hydrobiol. 27, 377-406. streams at higher altitudes, there was a conside Cummins, K.W. & Klug, MJ. 1979. Feeding ecology rable decline in numbers of specimens of mayf of stream invertebrates. Ann. Rev. Ecol. Syst. /0, lies at higher altitudes. Laboratory studies have 147-172. demonstrated that mayfly nymphs can feed on Dahi, E. 1957. Rondane, mountain vegetation in algae only (McCullough et al. 1979) and there South Norway and its relation to the environ fore this feature ofdistribution may be explained ment. Skr. norske Vidensk. Akad. Oslo. I. mat. na more by a decrease in temperature than reduced turv. kl. /956-3,1-375. input ofterrestrial organic matter. A decrease in Egglishaw, H. 1969. The distribution of benthic in vertebrates on substrata in fast-flowing streams. J. temperature can influence both the length oflife Anim. Ecol. 38, 19-33. cycle of several species and the composition of Elliott, J.M. 1965. Invertebrate drift in a mountain mayfly communities (Brittain 1975, 1976, Sto stream in Norway. Norsk ent. Tidsskr. 13. neburner 1977, Humpesch 1979). According to 97-99. Dahl (I957) the mean temperature in the Dov Frost, S., Huni, A. & Kershaw, W.E. 1971. Evalua refjell mountains in June-August decreases by tion of a kicking technique for sampling stream 0.7°C per 100 m elevation. bottom fauna. Can. J. Zool. 49,167-173: Temperature seemed to be the most important Gjrerevoll, O. 1975. Vegetasjon og flora. In Gjrere voll, O. & Ryvarden, L. (Eds.) Norges nasjonal factor determining the distribution ofmayflies in parker, Dovrejjell. Ormtjernkampen. Forlag, Sted, the study area. However, the abrupt shift in the pp. 41-70. food habits above the tree line seemed to be rela Gledhill, C. 1960. The Ephemeroptera, Plecoptera ted to the reduced input of terrestrial organic and Trichoptera caught by emergence traps in matter and the high nutritive values of stream two streams during 1958. Hydrobiol. /5, algae. 179-188.

104 Humpesch, U.H. 1979. Life cycle and growth rates - 1983. Hydrografi og ferskvannsevertebrater i Ra of Baetis spp. (Ephemeroptera: Baetidae) in the la umavassdraget 1982. K. norske Vidensk. Selsk. boratory and in two stony streams in Austria. Mus. Rapport Zoo!' Ser. 1983-2,1-74. Freshwat. Bioi. 9, 467-479. Rabeni, C.F. & Minshall, G.W. 1977. Factors affec Koksvik, J.1. 1979. Ferskvannsbiologiske og hydrog ting rnicrodistribution of stream benthic insects. rafiske unders0kelser i Saltfjell-/Svartisornradet. Oikos 29, 33-43. Del VI. Oppsummering og vurderinger. K. norske Raddum, G. & Fjellheim, A. 1982. Dyr som lager for Vidensk. Selsk. Mus. Rapport Zool. Ser. 1979-4, milj0inforrnasjon. In: Nicholls, M. (Ed.) Vass 1-79. dragsovervdking og vannjorskning. Foredrag fra Larsen, R. 1968. The life cycle of Ephemeroptera in konferanse pa Finse 13-14.9.1982. Norsk Lim the lower part of Aurland river in Sogn and Fjor nologforening, pp. 92-101. dane, Western Norway. Norsk ent. Tidsskr. 15, Shapas, TJ. & HilsenhofT, W.L. 1976. Feeding habits 49-59. of Wisconsin's predominant lotic Plecoptera, Ep LilIehammer, A. 1984. Distribution, seasonal abun hemeroptera and Trichoptera. Great Lakes Ent. 9, dance and emergence of stoneflies (Plecoptera) in 175-188. the 0vre Heimdal area ofthe Norwegian Jotunhe Steine, I. 1972. The number and size of driftiljlg imen Mountains. Fauna norv. Ser. B. 31, 1-17. nymphs of Ephemeroptera, Chironomidae and -& Brittain, J.E. 1978. The invertebrate fauna of Sirnuliidae by day and night in the River Stranda, the streams in 0vre Heimdalen. Holarct. Ecol. 1, Western Norway. Norsk ent. Tidsskr. 19, 271-276. 127-131. Macan, T.T. 1980. The occurrence of Baetis rhodani Stoneburner, D.L. 1977. Preliminary observations of (Ephemeroptera) in the River Lune. Aquatic In the aquatic insects of the Smoky Mountains: alti sects 2, 171-175. tudinal zonation in the spring. Hydrobiol. 56, McCullough, D.A., Minshall, G.W. & Cushing, C.E. 137-143. 1979. Bioenergetics of a stream «collectof» orga Ulfstrand, S. 1975. Diversity and some other para nism, Trichorythodes minutus (Insecta: Epheme meters ofEphemeroptera and Plecoptera commu roptera). Limnol. Oceanogr. 24, 45-58. nities in suharctic running waters. Arch. Hydro Minshall, G. & Kuehne, R. 1969. An ecological study bioi. 76, 499-520. ofinvertebrates ofthe Duddon, an English moun Vannote, R.L., Minshall, G.W., Curnmins, K.W., Se tain stream. Arch. Hydrobiol. 66, 169-191. dell, J.R. & Cushing, C.E. 1980. The river conti Nordhagen, R. 1943. Sikilsdalen og Norges fjellbei nuum concept. Can. J. Fish. Aquat. Sci. 37, ter. Bergens Mus. Skr. 22, 1-67. 130-137. N6st, T. 1981. Ferskvannsbiologiske og hydrografi Wooda1l, W.R. & Wallace, J.B. 1972. The benthic ske underS0kelser i Drivavassdraget 1979 - 80. K. fauna in four small southern Appalachian norske Vidensk. Selsk. Mus. Rapport Zool. Ser. streams. Am. Midi. Nat. 88, 393-407. 1981-/0,1-77. Received 4 March 1985

105 has been reported from floodrefuse and in moist, rot ( Short communications ) ting alder tree (Alnus) (Rozkosny 1973). ACKNOWLEDGEMENTS ODONTOMYIA ARGENTATA (FABRICIUS, 1794) (D/PT.. STRATIOMYIDAE) NEW TO We are indepted to Dr. R. Rozkosny, Department of THE NORWEGIAN FAUNA Environmental Studies, Faculty of Science, J.E. Pur kyne University, 61137 Brno, Kotlai"ska 2, Czechos LITA GREVE AND FRED MIDTGAARD lovakia, for verifying the determination of the speci Odontomyia argentata (Fabricius, 1794) is reported men of O. argentata. We are grateful to the Ministry new to the Norwegian fauna, and a short description of Environment for fmancial support. of the locality is given.

Uta Greve, Zoological Museum, University of Ber References gen, Museplass 3, N-5000 Bergen, Norway. Hulten, E. 1971. Atlas over vaxternas utbredning i Fred Midtgaard, Norwegian Forest Research Insti Norden. Generalstabens litografiska Anstalts For tute, P.O.Box 61, N-1432 As-NLH, Norway. lag, Stockholm: I- 531. In connection with a study of the insect fauna of Lundbeck, W. 1907. Diptera Danica. Vo!. 1. Stratio Ost0ya in the Oslofjord. a large material ofseveral or myiidae, etc. Pp. 1-75. (Stratiomyidae). Copen ders has been collected. Light-traps, Malaise-traps hagen. and window-traps have been used through-out the Rozkosny, R. 1973. The Stratiomyioidea (Diptera) of summer seasons 1983 and -84. In addition some ma Fennoscandia and Denmark. Fauna ent: Scand. terial were collected with insect-nets on visits of the 1.1-140. area. - 1982, 1983. A Biosystematic Study of the Euro pean Stratiomyidae (Diptera).1 & 1~ Dr. W. Junk On AK Brerum: Ost0ya (EIS:28) one female ofOdon Pub!. The Hague - Boston - London, 401, 431 tomyia argentata (Fabricius, 1794) was collected with pp. a net on 31 May 1984. The specimen was caught sweep-netting on a small grassy meadow, near Oust Received 16 Oct. 1984 gard, surrounded by a rich vegetation ofsmaller trees and herbs. OS10ya was also visited on 2 June, but no more specimens were found. O. argentata has not earlier been reported from BAET/SD/GITATUS BENGTSSON Norway (Rozkosny 1973). The species seems rare, (EPHEMEROPTERA), A NEW MAYFLY but widespread in Sweden northwards to Uppland and Varmland (Rozkosny 1982). Only two specimens SPECIES FOR NORWAY are known from Finland, the Helsingfors area, and in JOHN E. BRITTAIN Denmark the species has been reported as very rare from Zealand, Lolland and Falster (Lundbeck 1907; Abstract Rozkosny 1982). Rozkosny (I982) also map the Eu The mayfly Baetis digitatus is recorded from Norway ropean distribution of the species. for the first time. The nymphs of this species have Among the six species of Odontomyia recorded been collected at several localities in the large low from Fennoscandia (Rozkosny 1973); only O. hydro land river, Drammenselva, and two ofits tributaries, leon (L.) and O. microleon (L,) are known from south Snarumselva and Bingselva, in south-eastern Nor eastern Norway. O. hydroleon has the abdomen mar way. ked with a pattern of black and yellow or green, O. microleon has a shining black abdomen with a light Dr. John E. Brittain, Zoological Museum, University yellow lateral markings. O. argentata has a dull dor of Oslo, Sarsgate I, 0562 Oslo 5, Norway. sal part of abdomen in both males and females. The male O. argentata has the dorsal abdomen covered In connection with pollution monitoring studies of with silvery hairs, the female has transverse stripes of the Drammenselva watercourse in south-eastern golden-grey adpressed hairs. The genitalia of all the Norway (59°50'N, 9°55'E) several nymphal speci three species are figured in Rozkosny (1973, 1982). mens ofthe mayfly Baetis digitatus Bengtsson, 1912, O. argentata is an early summer species and there were taken both in the main river, Drammenselva, fore might have been overlooked. Lundbeck (I 907) and in two tributary rivers, Snarumselva and Bings reports the female to be sluggish while the males elva. were fast fliers. The species has been collected on Be The main river, Dramrnenselva, has a total catc rberis. Crataegus and Salix cinerea (Rozkosny 1982). hment area of 17 096 km2 and a mean discharge of Berberis vulgaris L. and Salix cinerea L. are com 330 m3 sec-I. The present localities are situated in mon on Ost0ya. S. cinerea has a distinct south-eas the lowest part of the catchment (Dramrnenselva) tern distribution in Fennoscandia (Hulten 197 I). The along a 46 km stretch of river which falls 63 m be two other plants are more widespread. The larvae fore it enters the Oslofjord at Drarnmen. Snarum-

106 Fauna norv. Ser. B, 32: Oslo 1985. selva and Bingselva have mean discharges of 110 and Almost half the Norwegian mayfly fauna, 19 spe 2 m 3 sec-I, respectively. All the localities have a cies, has been recorded from Drammenselva and its moderately strong current and a stony substratum, two tributary rivers Snarumselva and Bingselva. This sometimes with patches of aquatic mosses. The fol area appears to be the western limit for a number of lowing values indicate the range in mean chemical mayfly species which have reached Norway from the composition at the localities: pH 6.7-7.0; conducti east. These include Hepatgenia joernensis (Bengt vity 1.9-3.8 ~S m-I; total phosphorous 6.5-19.5 sson), Metretopus borealis (Eaton) and Ephemerella )JgPI-I. mueronata (Bengtsson). Baetis digitatus may well be The exact localities, and the number of nymphs long to this faunal group, although its occurrence in collected are: Drammenselva at Vikersund (UTM south-western Sweden and Denmark make this un ref.: NM557480, 6 nymphs; Drammenselva below certain. Embretsfoss (UTM re£.: NM518409), 57 nymphs; Drammenselva below D0vikfoss (UTM ref.: NM51039 2), 3 nymphs; Drammenselvaat Hokksund References (UTM ref.: NM509270), 2 nymphs; Drammenselva Belfiore, C. 1979. Segnalazione di Baetis digitatus Be at Nedre Eiker (UTM re£.: NM594248), 2 nymphs; ngtsson in Italia (Ephemeroptera, Baetidae). Bol/. Snarumselva (UTM ref.: NM527446), 13 nymphs Ass. Romana entomol. 34, 23-25. and Bingselva (UTM ref.: NM49234J), 78 nymphs. Bengtsson, S. 1912. Neue Ephemeriden aus Schwe The nymphs were collected in the months of April, den. Entomol. Tidskr. 33,107-117. May, July, October, November and December du Clifford, H.F. 1982. Life cycles ofmayflies (Epheme ring 1982, 1983 and 1984. roptera), with special reference to voltinism. Qu B. digilatus is closely related to B. niger and both aest. Entomol. 18, 15-90. species were recorded from several ofthe above loca Jensen, C.F. 1984. De danske Baetis-arter (Epheme lities. However, the hind margin of the last nymphal roptera: Baetidae). Flora og Fauna 90, 97-102. gill of B. digilatus is concave in contrast to that of B. KelTermiiller, M. 1978. Badania nad faunajetek (Ep niger which is convex. There are also other differen hemeroptera) Wielkopolski. VI. Badania Fizjog ces, such as the number of bristles on the tip of the raficzne nad Polska Zachodnia, Ser. C Zool. 31, glossa and in the antennae and cerci which can be 96-103. used to distinguish the two species (Miiller-Liebenau Miiller-Liebenau, I. 1969. Revision der europiiischen 1969). The two lighter patches on the abdominal Arten der Gattung Baetis Leach, I 815 (Insecta, terga are also characteristic in older nymphs. The Ephemeroptera). Gewass. Abwiiss. 48/49, adults are more difficult to separate (Miiller-Liebenau 1-214. 1969). Miiller-Liebenau, I. 1974. Baetidae aus Siidfrankre B. digitatus has a wide, but very patchy and gene ich, Spanien und Portugal (Insecta, Ephemeropte rally eastern distribution throughout much ofEurope ra). Gewiiss. Abwass. 53/54, 7-42. from the Balkans to Fennoscandia (Miiller-Liebenau Putz, V. 1978. Ephemeroptera. In: J. llIies (ed.) Um 1969, Putz, 1978). It was initially described from cen noJauna Europaea. Fischer, Stuttgart, pp. tral Sweden early this century (Bengtsson 1912). Sub 256-263. sequent records include Austria and England (in Savolainen, E. & Saaristo, M.I. 1980. Baetis digilatus Muller-Liebenau 1969), France (Muller-Liebenau Bengtsson (Ephemeroptera) found in Finland. No 1974) Poland (sowa 1975, KeffermUller 1978), Italy tul. Ent. 60: 195-196. (Belfiore 1979), Finland (Savolainenen & Saaristo Sowa, R. 1975. Ecology and biogeography of mayf 1980) and Denmark (Jensen 1984). lies (Ephemeroptera) of running waters in the Po On acCount of its rarity, B. digitatus is a poorly lish part of the Carpathians. I. Distribution and known species, athough it has been collected in a quantitative analysis. Acta Hydrobiol. 17: range of habitats. It has been recorded from slow 223-297. flowing small streams and rivers rich in aquatic vege tation (Muller-Liebenau 1969, Sowa 1975) and from Received 13 March 1985 the sheltered parts of fast-flowing rivers (Savolainen & Saaristo 1980). In Denmark and Sweden it has been found in the lower parts of large rivers (Jensen MACHIMUS INTERMEDIUS HOLMGREN IN 1984). Its disappearance from known localities in ZETTERSTEDT, 1852 (DIPTERA. ASILIDAE) Denmark during the last 20 - 30 years indicates its NEW TO NORWAY susceptibility to human activity. FRED MIDTGAARD The wide range in nymphal size at any given time within the Norwegian material suggests a long emer Machimus intermedius Holmgren in Zetterstedt, gence period and/or overlapping cohorts, which is 1852 is reported new to the Norwegian fauna, and a typical of many Baetis spccies (Clifford 1982). short review of its present status, and distribution is Nymphs were also recorded throughout most of the given. year, although from the few mature nymphs collec ted the emergence period is probably between May Fred Midtgaard, Norwegian Forest Research Insti and August. tute, P.O. Box 61, N-1432 As-NLH.

Fauntl norv. Ser. B. 32: Oslo 1985. 107 In the collection of Asilidae in the Zoological Mu synanthrophic beetle was introduced to Europe in seum of Oslo a female of Machimus intermedius 1961, probably from Africa, and has later appeared Holmgren in Zetterstedt, 1852 was found. The speci in several North Western European cities. The spe men was labelled: «Kristiania, Esmark, Machimus cies is known to feed on wool, fur, museum material, intermedius Z. det. Soot-Ryen 1926 Q». This species etc., and has become increasingly common in Scan has not formerly been reported from Norway. dinavia during the last twenty years. It was synomynized with Machimus setibarbus Loew, 1849 by Lyneborg (1968). M. setibarbus was Preben S. Ottesen, Department of Biology, Division assumed to have a very wide distribution from Scan of Zoology, University of Oslo, P.O. Box 1050, dinavia to the Mediterranean and east to Middle Asia. Blindern N-0316 Oslo 3. Examination of the genitalia of M. intermedius and specimens from various localities in the Mediterra In 1961 a brown and golden hairy Attagenus sp., nean showed that they differ distinctly from the geni 2.5 to 4.0 mm long, suddenly appeared in Moscow, tilia ofM. setibarbus from the type locality (Asia Mi and later in several cities in North Western Europe. nor). There are apparently several species which The systematic position of the species was much de have been identified as M. setibarbus in the past (The bated until Zhantiev (1973) finally proved it to be odor 1980). new to science and named it Attagenus smirnovi. It Because of the present uncertainty concerning the was found to interbreed with beetles collected on status of the taxa the best will be to use Holmgren's farms in Kenya, where African swift nests are name to the Scandinavian specimens (Lyneborg, thought to be its original habitat. Various synonyms pers. corn.). are treated by Arevad (1975) and Peacock (1979), Machimus intermedius was only known from who also give more detailed descriptions ofthe spe Sweden, Finland and USSR. In Sweden it has been cies, the latter including larvae. Full-size drawings found in Ostergotland, SOdermanland, bland and are found in Keiding (1970) and Hagstrom (1981). Skane. From Finland a few specimens are known The first specimen of A. smirnovi id Western Eu from the southern and eastern parts, and in USSR rope was caught in Gothenburg, Sweden in 1962. It one specimen has been recorded from the Leningrad was published as A. piceus Oliver (Hagstrom 1962, area (Lyneborg, 1968). The Norwegian fmd is the 1972; Hagstrom & Tornvall 1969), but later corrected first outside the Baltic area. (Hagstrom 1981). It appeared in Copenhagen 1963 Little is known about the biology, but Melin (I 923) (Hansen 1965) and London 1978 (Peacock 1979). reports one fmd from a little gravel pit with small On I Nov. 1983 one specimen was caught on a birches, ferns and grass, in the neighbourhood of a curtain in a student dormitory at Majorstua, Oslo by mixed coniferous forest. Ingar Iversen. The species has not previously been recorded from Norway (Silfverberg 1979). A. smirnovi is one of several foreign Dermestid be ACKNOWLEDGEMENTS etles which have recently invaded Europe. In Africa, I am greatly indebted to Dr. Leif Lyneborg for infor which is thought to be the species' continent of ori mations and for verifying the identification, and to gin, it is now found in farm stores, warehouses, and the staff at the Zoological Museum ofOslo for loan of in stored products. In Europe it is more restricted to their collection of Asilidae. human habitation, and has become a serious pest in the USSR, where the larvae feed on materials of REFERENCES animal origin, mostly those containing keratin, like wool, wool products, furs, hides, feathers, etc. (Pea-' Lyneborg, L. 1968. Notes on two species of Machi cock 1979). Wool and fur damage is recorded in mus loew in Northern Europe (Diptera, Asilidae). Sweden, where also damage to museum collections Notut. ent. 48, 131-135. has been noted (Hagstrom 1972). However, no such Melin, D. 1923. Contribution to the Knowledge of damage has yet been registered in Denmark, where the Biology, Metamorphosis and Distribution of the larvae seem to be omnivorous, although dead in the Swedish Asilids. Zoot. Bidr. Upps. 8, 1-318. sects and bird seeds are preferred. In that country the Theodor, O. 1980. Diptera, Asilidae, Fauna Palaes species is usually found in bedrooms and kitchens of tina. Insecta 11. Jerusalem. private homes, often in clean, modern apartments, and imago may be found at any time ofthe year (A Received 2 Nov. 1984. revad, pers. comm.). Imago does not feed (Zhantiev 1976). In Denmark the species today has become firmly AITAGENUS SMIRNOVI ZHANTIEV (COL., established in urban districts, especially in the Copen DERMESTIDAE) NEW TO NORWAY -A hagen area, but since 1977 also in some other towns. COMING INSECT PEST? Danish Pest Infestation Laboratory now receives PREBEN S. OTTESEN about 70 recordings per year which are with certa inty due to A. smimovi. No other species ofAttagenus On specimen of Attagenus smimovi Zhantiev was is reported more frequently from private homes (Are found on I. Nov. 1983 in an apartment in Oslo. This vad, pers. comm.). In Sweden it has been found in

108 Fauna norv. Ser. B. 32, Oslo 1985. eight districts from SkAne to Norrbotten (Palm 1982). was observed on young pines in Torshavn. This The species may thus be expected to spread in Nor dense larval population ofZ. diniana was parasitized way in the years to come. by the ichneumonid wasp Scambus eucosmidarum (Perkins). ACKNOWLEDGEMENTS Seppo Koponen, Zoological Museum, Department of I wish to thank Stig Lundberg, Lulea, Sweden, for Biology, University of Turku, SF·20500 Turku, Fin identifying the species, Kristian Arevad, Danish Pest land. Infestation Laboratory, Lyngby, for permission to include unpublished Danish observations, and Ingar INTRODUCTION Iversen, Univ. of Oslo, who found the fust Nor wegian specimen and asked me to identify and The vegetation of the Faroe Islands is dominated by publish it. grass heaths; no natural trees or higher shrubs grow there. Several plantations of coniferous and decidu ous trees have been established on the islands <0dum REFERENCES 1979, Hansen & 0dum 1982). One of the most com Arevad, K. 1975. Attagenus smirnovi Zhantiev (Cole monly used and best growing conifers is the lodge optera, Dermestidae): optrreden i Danmark. Ent. pole pine, Pinus contorta Douglas (Flensborg 1947, Meddr 43,172-176. Hansen & 0dum 1982). Hagstrom, T. 1962. Coleopterologiska meddelanden. In July 1984 I visited the Faroe Islands to study in Opusc. ent. 27, 59-60. sect herbivores in the plantations, and attacks by tort - 1972. Attagenus piceus - en inomhuslevande an ricid larvae on the lodgepole pines are presented ger pii spridning i Sverige. Fauna och Flora 67, here. 178-180. - 1981. Den bruna piilsangerns ratta identitet. RESULTS Fauna och Flora 76, 141-142. Feeding marks, larvae and pupae of Zeiraphera dini -& Tomvall, A.H. 1969. Meddelanden om coleop ana (Guem:e) were found on shoots of P. contorta in terfynd. Opusc. ent. 34, 133-135. late July on three Faroese islands: Streymoy, Eystu- Hansen, V. 1965. Nye danske biller. 1964. (Coleopte ra). Ent. Meddr 34, 123 -124. Keiding, J. 1970. Byer og huse som levested. In: N0r revang, A. & Meyer, TJ. (eds.) Danmarks natur 9. Det bebyggede land. Politiken. Copenhagen. pp. 346-371. Palm, T. 1982. Forandringar i den svenska skalbagg sfaunan. Ent. Tidskr. 103, 25-32. Peacock, E.R. 1979. Attagenus smirnovi Zhantiev (Coleoptera: Derrnestidae) a species new to Bri tain, with keys to the adults and the larvae ofBri tish Attagenus. Entomol. Gaz. 30, 131 -136. Silfverberg, H. (ed.) 1979. Enumeratio Coleopterorum Fennoscafzdia et Dania. Helsingfors Entomologi ska BytesfOrening, Helsingfors. Zhantiev, R.D. 1973. New and little known Dermes tids '(Coleoptera) in the fauna of the USSR. Zool. Zhurn. 52, 282 - 284. - 1976. Hide beetles (Family Dermestidae) of the fauna ofthe U.S.S.R. (In Russian). Moscow. 182 pp.

Received 2 Nov. 1984.

ZEIRAPHERA DIN/ANA (GUENEE) (LEP., TORTRICIDAE) ATTACKING PINUS CONTORTA DOUGLAS IN PLANTATIONS IN

THE FAROE ISLANDS • ",y ~-"""'--" SEPPO KOPONEN Larvae ofthe tortricid moth Zeiraphera diniana (Gu enee) were found to feed on planted lodgepole pines Fig. I. Occurrence of Zeiraphera diniana on Pinus on three islands of the Faroes. The heaviest damage contorta in the Faroes, 1984.

FaulUl norv. Ser. D, 32: Oslo 1985. 109 roy and Kunoy (Fig. 1). The heaviest damage was ob Faroes is Britain where the species feeds on lodgepole served on young lodgepole pines in a plantation near pines. the Faroese Academy at Debesartroo in Torshavn. The occurrence ofparasitoids in the dense Zeirap The average number of Zeiraphera larvae on two hem population in T6rshavn indicates a permanent metre tall pines was estimated to be about 100. Ho and probably long·term colonization of the tortricid wever, a marked proportion ofthese were dead or in in the Faroe Islands. P. contorta has been planted on poor condition, mainly due to parasitizing by ichneu a large scale on the islands since the 1920s especially monid wasps. On trees with a high density of larvae, after the year 1928 (S0ren 0dum pers. comm., FIens· young green cones were also damaged by them. borg 1947). The ichneumonid parasitoid found, High densities of Zeiraphera larvae were observed Scambus eucosmidarum (Perkins), is known as a pa locally on older lodgepole pines, one hundred metres rasite of some tortricids, including Z. diniana, and its from the above plantation. In Torshavn, Zeiraphera range includes hitherto the British Isles, Sweden, larvae also fed young lodgepole pines in plantations Germany, Switzerland and France (Aubert 1969). situated on a slope just below Hotel F0royar and in private gardens. No damage was seen in two old and large plantations of Torshavn: Gundadal (or the city park) and H0ydalUr. ACKNOWLEDGEMENTS Larvae of Z. diniana were observed in two planta tions on Eysturoy, Selatrad and S0ldarfj0rdur, and I wish to thank Dorete Bloch, Leivur Hansen and also in the remote plantation on Kunoy. Both young Trondur Leivsson (Torshavn) and S0ren 0dum and older lodgepole pines were attacked. Unfortuna (H0rsholm) for valuable information, Heli Hurme tely, the possible occurrence of Z. diniana in planta (Turku) for help in field work and Reijo Jussila (Tur tions on the two large islands in the south, Sandoy ku) for identifying the ichneumonids. I am indebted and Suduroy, was not investigated. to the Faroese Academy (Frodskaparsetur F0roya) In rearing experiments, the first adults of Z. dini for accomodation and laboratory facilities in Tors ana emerged on July 29, and the parasitic wasps havn. Financial support from the Nordiska Forskar about one week later. kurser and Kulturfonden for Finland och Danmark is acknowledged.

DISCUSSION REFERENCES Attacks by moth larvae on P. contorta in the Faroese Annila, E., Heliovaara, K., Puukko, K. & Rousi, M. plantations have been observed earlier (Trondur 1983. Pests on lodgepole pine (Pinus contorta) in Leivsson, pers. comm.). However, the identity of the Finland). Commun. Inst. For. Fenn. 115, 1- 27. pest has been unknown. The present pest species, Z. Aubert, J-F. 1969. Les lchneumonides Guest-Pa diniana, was recorded in the Faroes as early as the learctiques et leurs Hotes 1. Quatre Feuilles Edi 1950s when a female was found on Eysturoy (Wolff teur, Paris, 302 pp. 1970). Z. diniana is a wellknown pest of the larch Bakke, A. 1969. Extremely low supercooling point in (Larix decidua Miller) in the Alps (e.g. Schwenke eggs of Zeiraphera diniana (Guenee) (Lepidop 1978). In other parts ofEurope it is known to feed on tera: Tortricidae). Norsk ent. Tidsskr. 16, 81-83. many conifers, such as Pinus contorta in Upland Bri Day, K. 1984. Phenology, polymorphism and insect tain (Day 1984) and the Scots pine (P. sylvestris L.) in plant relationships of the larch budmoth, Zeirap Fennoscandia. The species has attacked heavily tree hera diniana (Guenee) (Lepidoptera: Tortricidae), line forests ofP. sylvestris in southern Norway (Bakke on alternative conifer hosts in Britain. Bull. ent. 1969). Contrary to the situation in Britain, Z. diniana Res. 74, 47-64. has not been found as a pest of planted P. contorta in Eidmann, H. H. 1982. Insekter pa contortatall i Sve Fennoscandia (Eidmann 1982, Annila et al. 1983, rige (Insects on lodgepole pine in Sweden). Sveri Karlman 1984). ges Skogsvdrdsforb. Tidskr. 1980: 1-2. 59-64. Larval colour varied greatly in the present mate Flensborg, C. E. 1947. PlantningsforS0gene paa Fae rial, the light type dominated over the dark one. Thus roorne. Hedeselsk. Tidsskr. 1947: 5-6.1-7. the situation resembled that on lodgepole pine in Bri Hansen, L. &0dum, S. 1982. Haverog plantager. In: tain (Day 1984). N0rrevang, A. & Lund0, J. (eds.), FaerfJrnes na According to WoltT (I971), Z. diniana is absent on tur, Politikens Forlag, K0benhavn, pp. 79-87. Orkney and Shetland (and Iceland), and a transport to Karlman, M. 1984. Pathogens and other threats of the Faroes by human agency, for example with tree Pinus contorta in northern Sweden. Univ. of seedlings, is very probable. At least some of the lod Umea, 212 pp. gepole pines near the Faroese Academy, attacked by Nyholm, I. 1970. Faer0rne - en uforglemmelig op Z. diniana, originated from Scotland and were plan levelse. Hedeselsk. Tidsskr. 1970: 14-15. ted in 1960 (Nyholm 1970). There has been no phy 300-305 & 320-327. tosanitary control over the transfer of plants from 0dum, S. 1979. Actual and potential tree-line in the Norway, Denmark, Scotland or Iceland to the Fa North Atlantic region, especially in Greenland roes. The most probably origin of Z. diniana in the and the Faroes. Holarct. £Col. 2. 222-227.

110 FaurlQ norv. Se,. B. 31, Oslo 1985. Schwenke, W. 1978. Die Forstschiidlinge Europas 3. logical Museum, University of Bergen. The localities Schmetter/inge. Verlag Paul Parey, Hamburg & are presented according to the system of 0kland Berlin, 467 pp. (I98 O. Wolff, N. L. 1970. Revidert fortegnelse over Faem ernes sommerfugle (A revised list of the Lepidop Tanypeza longimana Fallen, 1820 tera known from the Faroe Islands). Ent. Meddr. Published records: AK, Oslo: Oslo I d I Q (ZMO), 38,3-14. Bekkelaget 30 June 1847 I d IQ (ZMO); Enebakk: Wollf, N. L. 1971. Lepidoptera. The Zoology ofIce Enebakk I d (ZMO). HES, Elverum: Grundset.1 d, land Ill: 45,1-193. I specimen (ZMO). OS, S0ndre Land: Sk0ien I Q Received 14 Jan. 1985. (ZMO). ON, Dovre: Dovre 6 August 1973 1 Q (ZMO). NTI, Levanger: Tynset (Zetterstedt in Iitt). New records: AK, Frogn: H3.0ya 27 June-22 July TANYPEZA LONGIMANA FALLEN. 1820 (DIPT.• 1984 2 d d (Malaise trap A), 22 July-18 August TANYPEZIDAE) AND STRONGYLOPTHALMYIA 1984 I Q (Malaise trap B) (ZMB). BV, AI: Torpo 10 USTULA TA (ZETIERSTEDT, 1847) DIPT., July 1982 IQ (ZMB). TEY, Kragem: Skat0Y 8 July STRONGYLOPHTHALMYIIDAE) IN NORWAY 1983 I Q (ZMB). HOY, Bergen: Eidsvag, Vollane 9 UTA GREVE AND AGE SIMONSEN July 1984 I Q (ZMB). The Norwegian records total eleven, older mate ABSTRACT rial included. T. longimana seems to be rare in Scan Strongylophthalmyia ustulata (Zetterstedt, 1847) is dinavia and Lyneborg (I 962) reports only six records reported new to Norway. Four specimens were from Denmark. It is, however. a common species in found, one female near Valand, Mandal West Agder Central Europe where it can be found in schrubbe province and three females at the island Ha0ya, ries and on leaves on decidious trees, especially Po Frogn, Akershus province. For the first time since pulus tremula L. and Betula spp. The localities are of the survey of Norwegian Diptera hlade by Siebke ten situated near brooks and brooklets. T. longimana (0. (I 877) new records are given from Norway for Ta has Qvick pers. comm.) been recorded in Sweden nypeza longimana Fallen, 1820. The distribution of from humid sites with rich vegetation ofbushes (Pru both species in Norway is mapped. nus padus) and trees (Quercus robur). The new records from Norway are mainly from Uta Greve and Age Simonsen, Zoological Museum, localities with mixed decidious forests. At trap A, Ha University of Bergen, N-5000, Norway. 0ya, the forest was dominated by Tilia cordata Mill., Ulmus glabra Huds. and Quercus robur L. There were also several old Populus tremula L. The area McAlpine et al. (I 98 I) splits the old family Tanypezi had been left untouched for many years and old and dae into Tanypezidae and Strongylophthalmyiidae. dead trees had not been removed. The ground vege The families are placed together with Psi/idae and tation was rich and varied. The area near trap B was Diopsidae in the superfamily Diopsoidea. Tanypezi of a similar type. dae and Strongylophthalmyiidae are Acalyptrate flies. Only two females, from Skat0y and Torpo, were The Strongylophthalmyiidae can be distinguished netted. The female from Vollane was caught in a from the Psilidae and the Tanypezidae in the follo window inside a house. Bordering to the garden of wing chara*ristics: Notopleuron is usually with the house was a mixed, open forest of Tilia. Ulmus, two bristles and anepisternum with outstanding Betula, Fagus and flex. bristles in addition to fine hairs. The ocelli are situa The Norwegian distribution is mapped on Fig. I ted far forward, with the anterior one about midway A. Strongylophthalmyia ustulata (Zetterstedt, 1847) between vertex and antennae (McAlpine et al. 198 I). New records: AK, Frogn: Ha0ya 3-16 June 1984 Tanypezidae may be mistaken for Micropezidae. IQ, 27 June-22 July 1984 IQ, 22 July-18 Au Tanypezidae are distinguished by their large eyes, gust 1984 I Q (All Malaise trap A) (ZMB). VAY, distinctly higher than long. In Norwegian Tanypezi Mandal: Marnarvegen near road to Valand 6- 22 dae the tarsus of last pair oflegs is longer than the ti July 1982 I Q (Malaise trap) (ZMB). bia - in Micropezidae the tarsus oflast pair oflegs is These are the first records ofS. ustulata from Nor shorter than the tibia. way. All specimens were found in material from Ma Tanypezidae and Strongylophthalmyiidae are dis laise traps. S. ustulata is reported both from Finland tributed in the Holarctic. Each family is represented and Sweden. by one species only in the Fennoscandian fauna; Ta S. ustulata at H3.0ya was found together with T. nypezidae with Tanypeza longimana Fallen, 1820, longimana. The locality near Mandal was a boggy Strongylophthalmyiidae with Strongylophthalmyia area relatively rich in herbs and surrounded by ustulata (Zetterstedt, 1847). Previous records from mixed forests. 0. Qvick (Pers. comm.) has found Norway are given by Zetterstedt (I 847) and Siebke both species together in Sweden. (I 877) for T. longimana only. The records ofthe species indicate perhaps humid The present paper includes all material in the Nor loving, termophilic species with preference for relati wegian zoological museums. Abbreviations: ZMO vely mineral-rich habitat types. Zoological Museum, University of Oslo, 2MB-Zoo- The Norwegian distribution is mappes in Fig. I B.

Fauna nom Ser. B, 32 .. Oslo 1985. 111 Fig. I. Records ofTanypezidae and Strongylophthal myiidae in Norway. A- Tanypeza longimana Fal len, 1820, B- Strongylophthalmyia ustulata (Zetter stOOt, 1847). Open circle - Zetterstedt.

ACKNOWLEDGEMENTS We are indepted to Fred Midtgaard, k; and Alf-Ja - 1973. Ordnung Diptera (Zweifiiigler). Handb. cob Nilsen, Hidrasund for permission to sort out ma Zool. 4 (2) 2/31. 1-337. terial from their Malaise traps, and to Agnete and Lyneborg, L. 1962. Danske acalyptrate fiuer. I. Ent. Knut Rognes for material from Buskerud. Our Med. 31, 249-264. thanks are also due to Ullmar Qvick, Eskilstuna for McAlpine, J. F. et al., 1981. A manual of Nearctic information on habitats for both species in Sweden. Diptera I, 674 pp. Siebke, H. 1877. Enumeralio /n.seclorum Norvegico rum. Fasciculum IV. Catalogum Dipterorum Continentem. A. W. Bmgger. Christiania, 255 pp. REFERENCES SteySkaI, G. S. 1974. Recent advances in the primary Griffiths, G. C. D. 1972. The phylogenetic classifica classification of the Diptera. Ann. ent. Soc. Am. tion of Diptera Cyclorrhapha, with special refe 67, 513-517. rence to the structure of male postabdomen. Dr. Zetterstedt,1. W. 1847. Diptera Scandinaviae dispo W. Junk, N. V., The Hague. 340 pp. sita et descripta. 6, Lundae, p. 2422. Hennig, W. 1937. Tanypezidae. In Lindner, E. (00.). 0k1and, K. A. 1981. Inndeling av Norge ill bruk ved Die Fliegen der Palaearktischen region 5 (44), biogeagrafiske oppgaver - et revidert Strand-sy 1-6. stem. Fauna, Oslo 34, 167-178.

112 Fauna 110",. Ser. B, 32: Oslo 1985. GUIDE TO AUTHORS. References. In the text: Black (] 979), Black & Blue (] 973: I00), or «as noted by Green (] 978) and Black FAUNA NORVEGICA Ser. B. publishes papers in (I 979)". Multiple references should be given in chro English, occasionally in Norwegian and German with an extensive English abstract. Contributors nological order, i.e. (Black & Blue 1973, Green 1976, 1979, Black 1978). with a native language other than the language used in the paper submitted, are requested to have manu List of references are to be unnumbered and in inter national alphabetical order (i.e. = AA, lE and scripts linguistically revised prior to submission. A When preparing manuscripts for submission, au A =Ae, 0 and 0 =Oe). Titles of journals should be abbreviated according to the World List of Scientific thors should consult current copies of Fauna norve Periodicals. Do not refer to papers (dn prep.» among gica and follow its style as closely as possible. the references. Manuscripts not conferring to the guide to authors will be returned for revision. Examples: Journal: Manuscripts should be submitted to the Editor-in Chief. Send two copies. They must be typewritten, L0ken,A. 1962. Social wasps in Norway (Hymenop tera, Vespidae). Norsk ent. Tidsskr. 12, 191 - 218., ~ double spaced throughout, on one side of the paper, and wide margins, 5-6 cm on the left. Separate Book: sheets should be used for the following: I) Title page, Mayr, E. 1913. Animal species and evolution. Har I with author's name. 2) An abstract, with the name vard University Press. Cambridge, Mass. and full postal address of the author underneath. 3) Fittkau,E.J. 1962. Die Tanypodinae (Diptera, Chiro Tables with their headings. 4) Legends to figures. nomidae). Die Tribus Anatopyniini, Macropelo Dates should be referred to as 10-20 Aug. 1970. poni und Pentaneurini. AM. Larvalsyst. Insekten Underline all generic and species names. Other 6,453 pp. underlinings should be left to the editor. Approxi Chapter: mate position of figures and tables in the text should be indicated in the margin. All acknowledgements Whitman,L. 1951. The arthropod vectors of yellow should be given under a single heading at the end of fever.- In: Strode,K. (ed.), Yellow Fever. Mc. the text, but before the references. Graw - Hill, New York & London, pp 229 - 298. Figures and Tables. Send two copies. All illustra tions should be identified lightly with the author's Proofs. Two copies of the first proof will be sent to name and the figure number. The figures and tables the author: One corrected copy should be returned to should be constructed in proportion to either the en the editor without delay. Alterations should be limi tire width of the typed area (140 mm) or to the co ted to correcting typesetting errors. Extensive altera lumn width (67 mm). tions will be charged to the author. Nomenclatlue. The first time a binomen is used in the text the name of its author should be included. Reprints. Twentyfive reprints are supplied free (fifty Author names should be written in full except L. for with multiple authorships). Additional reprints can Linneaus. Dates can be included when considered be ordered at a charge (an order form is sent with the necessary, i.e. 'Rhyacophila nubila (Zetterstedt, 1840). proofs).

FAUNA NORVEGICA Ser. A, B, C vii normalt ut Referansemessig skalen aldri ta hensyn tit numme komme med til sammen 5 hefter i 10pet av en argang ret i IiJvre hliJyre hjliJrne pd omslaget Gnne i firkanten). (eH volum). For at heftene skal komme inn undet Det en skal ta hensyn til er de oppgitte data for re Postverkets regler for billig serie-utsendelse, for spektive serier. De f0rste heftene i hver serie vii i ar langes det at Fauna norvegica i hvert kalenderar gis gang 1981 hete henholdsvis Ser. B'vol. 28 No. I. Ser. fortl0pende nummer fra I og oppover (altsa i argang C Cinclus Vo!. 4 No. I, Ser. A Vo!. 2 No. I. Det er 1981: 1-5). Det vii kunne bli noe tilfeldig hvilke disse data sam gir den korrekte Iitteraturreferansen, hefter som blir gitt de respektive nummer pa grunn og det er disse forkortelsene som star oppf0rt i Ab av uregelmessigheter med rekkefelgen i 10pet av aret. stract til hver artikkel og pa srertrykkene.

, 9 S2 S8 or· 62 80 83 89 94 97 100

OdoruomJlUl argenlillQ (Fabricius. 1794) (Oipt., Stratiomyidae) n, to the 106 106 107 10 109 III