Norwegian Journal of Entomology
Volume 47 No. 2 • 2000
Published by the Norwegian Entomological Society Oslo and Stavanger NORWEGIAN JOURNAL OF ENTOMOLOGY
A continuation of Fauna Norvegica Serie B (1979-1998), Norwegian Journal ofEntomology (1975 1978) and Norsk Entomologisk Tidsskrift (1921-1974). Published by The Norwegian Entomological Society (Norsk entomologisk forening).
Norwegian Journal of Entomology publishes original papers and reviews on taxonomy, faunistics, zoogeography, general and applied ecology of insects and related terrestrial arthropods. Short com munications, e.g. less than two printed pages, are also considered. Manuscripts should be sent to the editor.
Editor Lauritz Sl/lmme, Department of Biology, University of Oslo, PO.Box 1050 Blindem, N-0316 Oslo, Norway. E-mail: [email protected].
Editorial secretary Lars Ove Hansen, Zoological Museum, University of Oslo, Sarsgate I, N-0562 Oslo. E-mail: [email protected].
Editorial board Ame C. Nilssen, Tromsl/l John O. Solem, Trondheim Lita Greve Jensen, Bergen Knut Rognes, Stavanger Ame Fjellberg, Tjl/lme
The goal of The Norwegian Entomological Society is to encourage the study of entomology in Norway and to provide a meeting place for those who are interested in the field. Annual membership fees are NOK 200 (juniors NOK 100) for members with addresses in Norway, and NOK 220 (Juniors NOK 110) for members abroad. Inquiries about membership should be sent to the secretary: Jan A. Stenll/lkk, P.O.Box 386, N-4002 Stavanger. Norway. E-mail: [email protected].
Norsk entomologisk forening (NEF) ser som sin oppgave a fremme det entomologiske studium i Norge, og danne et bindeledd mellom de interesserte. Medlemskontingenten er kr. 200 (junior kr. 100) pr. ar for innenlandsmedlemmer og kr 220 (junior kr. 110) for medlemmer bosalt utenfor Norge. Forespl/lrsel om medlemsskap i NEF sendes sekret NEF web site: http://www.entomologi.no/ Subscription. Members of The Norwegian Entomological Society receive Norwegian Journal of Entomology and Insekt-Nytt free. Institutional and non-member subscription NOK 250. Subscriptipn and membership fees should be sent to the treasurer: Bjl/lm 0kland, Brekkeskog 31, N-1430 As, Norway; account no.: 08065440920. Front cover: Boreophilus henningianus Sahlberg, 1832 (Coleoptera, Staphylinidae). Artist: Frode 0degaard. Printed by: Reprografisk Industri AS, Oslo. ISSN 1501-8415. Norw. J. EntomoJ. 47, 101-112.2000 Diel fluctuations of invertebrate drift in a Norwegian stream north of the Arctic Circle Morten Johansen, J. Malcolm Elliott & Anders Klemetsen Johansen, M., Elliott, J.M. & Klemetsen, A. 2000. Die1 fluctuations of invertebrate drift in a Norwe gian stream north of the Arctic Circle. Norw. J. Entomol. 47, 101-112. Sreterelva is a small, third order, stream situated north of the Arctic Circle (latitude 68° N). Drift samples were collected in 1996 during four periods (22-24 May, 23-25 June, 18-20 August and 6-7 October) to determine whether diel fluctuations in drift changed with the changing light regimes of the Arctic summer. Two of the sampling periods (May and June) had continuous daylight, while August and October had dark nights. The taxonomic composition ofthe drift in Sreterelva was similar to that ofother northern temperate streams, being dominated by larvae of Ephemeroptera, Plecoptera and Chironomidae. Total 24 h drift density (± SE) was high, ranging from 242 ± 45 individuals per 100 m) in May to 772 ± 66 per 100 rn' in October. Terrestrial invertebrates contributed a high proportion to the total drift (7-39 %), probably as a result ofthe dense riparian vegetation along the stream. Total drift density of aquatic invertebrates was greater at night than during the day in August and October, while there were no differences between day and night in May and June. Most individual taxa of aquatic invertebrates followed this pattern. The major exception was Hydracarina, which consistently drifted with highest density during the day. It is concluded that simple classifications of drift are difficult to apply to Arctic streams with periods of continuous daylight. Key words: invertebrate drift, aquatic invertebrates, die1 fluctuations, streams. Morten Johansen & Anders Klemetsen, Department ofMarine and Freshwater Biology, University ofTromso, N-9037 Tromso, Non1/ay. J. Malcolm Elliott, Freshwater Biological Association, The Ferry House, Far Sawrey, Ambleside, Cumbria LA22 aLp, United Kingdom. INTRODUCTION causes have been studied extensively (reviews by Waters 1969, 1972, Muller 1974, Statzner et al. Rivers and streams differ from other aquatic en 1984, Brittain & Eikeland 1988, Allan 1995). vironments in having a strong unidirectional flow of water that transports materials from upstream Elliott (l965a) published the first account of in to downstream areas. This transport forms the ba vertebrate drift in Norway. Since then, there have sis of the River Continuum Concept (Vannote et been relatively few publications on this phenome al. 1980, Minshall et al. 1985). Physical forces non in Norwegian streams (Steine 1972, Fjellheim associated with the current are perhaps the most 1980, Brekken et al. 1981, Sandlund 1982, Lil important factor affecting organisms in running lehammer et al. 1995). Invertebrate drift was in water and many invertebrates possess morphologi vestigated over several years in Kaltisjokk, a cal adaptations which help them avoid being swept Swedish stream just north of the Arctic Circle (66 away. Nevertheless, many of these organisms en ON) (Muller 1966, 1970). The present account is ter the water column (either actively or passively) the first on invertebrate drift in a Norwegian stream and are displaced downstream by the current. This north of the Arctic Circle. phenomenon (termed invertebrate drift) and its 101 Johansen et al.: Diel fluctuations of invertebrate drift in a Norwegian stream Since Tanaka (1960) described die1variations in S~terelva is an important spawning site for brown the composition and the intensity ofthe drift, simi trout (Salmo trutta L.), and the fish community lar patterns have been observed frequently all over consists mainly ofjuvenile trout with a few three the world (e.g. Waters 1962, Elliott 1965a, b, 1967, spined sticklebacks (Gasterosteus aculeatus L.). McLay 1968, Bishop & Hynes 1969, Clifford 1972, Hynes 1975, Cowell & Carew 1976, Bailey METHODS 1981, Allan et al. 1988, Brewin & Ormerod 1994). Several studies have shown that light is the major The drift of invertebrates was sampled over two cue for the diel periodicity ofthe behavioural drift nights and one day during each of three periods: (e.g. Muller 1965, Elliott 1967, Holt & Waters 22-24 May, 23-25 June, 18-20 August 1996. One 1967, Pearson & Franklin 1968, Bishop 1969, day and one night was also sampled on 6-7 Octo CoweIl & Carew 1976). North of the Arctic Cir ber 1996. Drift was sampled using four floating cle, daylight is continuous in summer, but progres drift nets with a mesh size of 430 Ilm (a detailed sively shortens in autumn to a winter minimum. description is given in Elliott 1967). The nets were emptied every 3 h. Each net sampled to a depth of The main objectives of this study were to exam 6 cm and the total submerged area was 288 cm'. ine how the composition and the dieI fluctuations An Ott miniature current meter was used to meas of the invertebrate drift changed with season. ure water velocity at the middle ofthe net mouth, at the beginning and end of each 3 h-sample. The STUDY SITE volume of water filtered was estimated from the product of the area of the submerged net mouth, S~terelva is a small third order stream which is current velocity and duration ofsampling. All drift the main tributary of the lake Mokkelandsvatn in samples were preserved in 70 % ethanol, and later southern Troms (northern Norway) north of the sorted in the laboratory using a dissecting micro Arctic Circle (68±49DN, l6±27DE) (Figure 1). scope at 6x magnification. The invertebrates were The stream flows through mixed deciduous for handpicked, identified and counted. ests, with riparian vegetation dominated by grey alder (Alnus incana (L.) Moench), birch (Betula Flow through the nets was used to calculate drift pubescens Ehrhart) and willow (Salix spp.). The density as the number ofdrifting invertebrates per catchment geology is dominated by micaceous 100 m3 sampled (Waters 1969, Elliott 1970, Al1an gneiss and slate. The flow regime is characterised & Russek 1985). Mean drift densities per samp by wide seasonal fluctuations, with peak flows ling date (referred to as 24 h drift density) were during snowmelt in May-June and the lowest flows calculated by pooling all samples from each net 3 during winter. The mean annual discharge is 0.5 m / and dividing by the total number of hours sam s. Water temperature falls below 1 DC in early No pled at that time. Before statistical analyses, all vember, and the stream is ice covered from No drift densities were log10 transformed to ensure vember to May. The area has a coastal climate, that the components of variance were additive with relatively mild winters and cool summers. before using analysis ofvariance (ANOVA) to test The stream is oligotrophic (total nitrogen 85-130 for temporal differences in drift (Elliott 1971). A flg/l, total phosphorus 2-6 flg/l) and circumneutral two-way ANOVA using the General Linear Model (mean pH 7). The stream is mildly polluted, mainly (GLM) was performed with day versus night as from sewage and agricultural runoff (Muladal & one main effect and sampling date as the other Skotvold 1993). main effect. Night in August and October was de fined as the period ofdarkness between sunset and A 200 m-long stretch of the stream just upstream sumise. Even though there was no darkness on from Mokkelandsvatn was chosen as the study the sampling occasions in May and June, there was reach. It has a stony substratum consisting of still a large diel difference in light intensity. To gravel, cobbles and a few boulders. Aquatic macro test statistically for a diel difference in drift caused phytes and bryophytes are absent. The mean chan by darkness, «night» in May and June was defined nel width is 4 m and the mean depth is 0.2 m. 102 Norw. J. Entomol. 47, 101-112.2000 as the two 3 h samples taken around midnight m3Is) and highest in May (1.6 m3Is). Discharge in (22:00 to 04:00). June and October was 0.7 and 0.8 m3/s, respec tively. RESULTS Physical measurements Drift composition Temperatures ranged from a low of2.4 QC in May Altogether, 91 taxa (of which 74 were aquatic in to a high of 13.1 QC in August. Diel variation was vertebrates) were collected and identified in the highest in August (range 9.9-13.1 QC), lowest in drift samples during the study. The highest number October (range 5.2-5.5 QC) and intermediate in ofaquatic taxa was collected in June (64), the least May and June (2.4-3.7 QC and 5.7-6.4 QC, respec in August (38). Emerging aquatic insects contribu tively). River discharge was lowest in August (0.1 ted on average 12 % (range 0-31 %) to total drift. N A Arctic Circle) f4~'" " Bergs vagen o 1 km Figure 1. Map of the stream Saaterelva with the study location marked. 103 Johansen et al.: Diel fluctuations of invertebrate drift in a Norwegian stream The terrestrial component of the drift (inverte by newly-emerged Chironomidae. Adult Plecopte brates with a completely non-aquatic life cycle, ra (mainly emerging Brachyptera risi (Morton)) such as some Hemiptera, Arachnida, Collembola drifted significantly more at «night» (F(2.m=5.08, and Coleoptera) contributed an average of 22 % P<0.05). Water mites drifted significantly more (range 7-39 %). during the day than at «night» (F(2.371=14.25, P 104 Norw. J. Entomol. 47, 101-112.2000 followed that for total aquatic invertebrates with There was no difference in the density of drifting the highest drift densities in the first half of the Chironomidae (both larvae and pupae) between night (Figure 2c). In October (Figure 3d), mean day and «night» in May (Table I). In June the high drift was significantly higher at night than during est density was found during the day, and the dif the day (F(1,3w4.35; P 1200 (a) 1000 <'"1"'-'" 8 800 o 600 o ..--;< 400 \-< (j) 200 0., 1200 (b) 20 0 1000 ~ ,,! [ 800 101l. 600 400 200 o+---+-~-~---~_~__-+---~ 01 04 07 10 13 16 19 22 01 04 01 04 07 10 13 16 19 22 01 04 Time of day C:J Aquatic invertebrates _ Emerging aquatic insects r::z2ZI Terrestrial invertebrates Figure 2. Stacked drift densities (number of individuals per 100 m3 for the three main drift components in Sceterelva. Changes in the water level during sampling are also given. Samples from (a) 22-24 May, (b) 23-25 June, (c) 18-20 August, and (d) 6-7 October 1996. Shaded bars on the x-axes represent period of darkness between sunset and sunrise. 105 Johansen et al.: Diel fluctuations of invertebrate drift in a Norwegian stream Table 1. Mean drift densities (number of individuals per 100 m3 ± 1 SE) of common invertebrates caught on the four sampling occasions in Sreterelva, northern Norway. SE < 0.5 not shown. +, taxa present; -, not found. *, P<0.05; **, P<0.01; *** P<0.001. P-values are from ANOVAs between the 3 periods in May, June and August, and the 2 periods in October. The «night»-period in May and June is the time of day equivalent to the period of darkness in August (termed night). May June «Night» Day «Night» «Night» Day «Night» Aquatic invertebrates Plecoptera 14 ± 2 17 ± 2 11 ± 2 21 ± 4 21 ± 1 22 ± 2 Brachyptera risi 4 ± 1 3 3 ± 1 9 ± 2 5 ± 1 10 ± 2 Leuctra spp. 2 3 ± 1 2 1 + + Nemouridae 7 ± 2 11 ± 1 5 ± 1 11 ± 2 14 ± 1 11 ± 1 Capnidae + 1 + + + + Ephemeroptera 20 ± 2 22 ± 2 17 ± 3 70 ± 16 61 ± 4 66 ± 9 Baetis spp. 20 ± 2 19 ± 2 17 ± 3 65 ± 16 54 ± 4 60 ± 9 Ephemerel/a aurivil/ii + + + + + + Diptera 173 ± 44 152 ± 26 79 ± 13 147 ± 14 193 ± 11 158 ± 13 * Chironomidae 38 ± 9 34 ± 4 26 ± 3 120 ± 12 159 ± 10 128 ± 12 * Simuliidae 2 ± 1 3 2 ± 1 23 ± 2 29 ± 2 25 ± 2 Tipulidae 4 ± 2 1 2 ± 1 * + + + Limonidae 1 1 1 + 1 + Empididae 3 ± 1 2 2 ± 1 2 2 2 Psychodidae 123 ± 38 110 ± 23 46 ± 10 1 1 1 Coleoptera 1 3 ± 1 1 21 ± 14 18 ± 9 24 ± 10 Trichoptera 1 1 1 2±1 2 2 Rhyacophila nubila 1 + + 1 1 1 Limnephilidae + + + + + + Hydracarina 3 ± 1 3 ± 1 2 ± 1 9 ± 1 26 ± 3 10 ± 1 Cyclopidae + + + 6±2 8 ± 2 7±3 Oligochaeta 13 ± 4 5 ± 1 9 ± 3 * 2 2 2 Total 227 ± 48 205 ± 31 121 ± 15 278 ± 41 332 ± 17 291 ± 22 Emerged aquatic insects Plecoptera + 2 ± 1 + 4 ± 1 2 4 ± 1 Ephemeroptera - - - - + Chironomidae + 3 ± 1 + * 66 ± 20 181 ± 29 57 ± 14 * Trichoptera - + - + + + Total 1 5 ± 2 + * 77 ± 22 193 ± 30 67 ± 17 * Terrestrial invertebrates Diptera 4±2 23 ± 12 3 ± 1 5±2 6 ± 1 5±2 Hemiptera 1 ± 1 15 ± 6 1 * 32 ± 3 43 ± 4 32 ± 5 Coleoptera + 2 ± 1 - 2 3 ± 1 1 Hymenoptera - + - 1 2 1 Lepidoptera - + + 1 1 1 Collembola 11 ± 3 28 ± 9 5 ± 2 17 ± 3 34 ± 6 18 ± 3 Total 17 ± 4 67 ± 24 10 ± 3 61 ± 6 92 ± 10 60 ± 9 Total, all groups 245 ± 52 277 ± 54 131 ± 17 416 ± 42 618 ± 43 418 ± 29 106 Norw. J. Entomol. 47, 101-112.2000 Table 1. Continued. August October Night Day Night Night Day Aquatic invertebrates Plecoptera 8±3 8 ± 1 44 ± 6 159 ± 16 52 ± 4 Brachyptera risi + 15 ± 2 16 ± 3 Leuctra spp. 1 3 4 ± 1 35 ± 5 8 ± 1 Nemouridae 5 ± 1 3 ± 1 30 ± 5 66 ± 7 26 ± 3 Capnidae 1 31 ± 3 5 ± 1 Ephemeroptera 13 ± 4 11 ± 1 26 ± 3 378 ± 54 103 ± 17 Baetis spp. 12 ± 3 10 ± 1 20 ± 3 371 ± 54 101 ± 17 Ephemerella aurivillii 1 + 6 ± 1 1 Diptera 55 ± 12 66 ± 8 325 ± 40 277 ± 27 141 ± 13 Chironomidae 33 ± 10 55 ± 7 238 ± 31 211 ± 24 98 ± 12 Simuliidae 19 ± 3 9 ± 1 65 ± 7 42 ± 4 19 ± 1 Tipulidae 1 1 7±2 1 + Limonidae + 9 ± 2 5 ± 1 Empididae + + 1 + Psychodidae + + + 7 ± 1 5 ± 1 Coleoptera 3 ± 1 2 20 ± 2 7 ± 1 7 ± 1 Trichoptera 1 4 14 ± 2 18 ± 2 10 ± 1 Rhyacophila nubila 1 1 2 1 1 Limnephilidae + 1 6 ± 1 16 ± 2 8 ± 1 Hydracarina 5 ± 1 15 ± 2 19 ± 2 11 ± 2 13 ± 1 Cyclopidae 2 ± 1 1 1 1 + Oligochaeta + + 1 7 ± 1 4 ± 1 Total 89 ± 18 111 ± 10 450 ± 51 859 ± 82 331 ± 33 Emerged aquatic insects Plecoptera + + + 2 Ephemeroptera 3 ± 1 + + + Chironomidae 56 ± 28 84 ± 13 84 ± 12 8 ± 2 25 ± 5 Trichoptera + + + Total 57 ± 25 90 ± 14 93 ± 12 11 ± 2 31 ± 6 Terrestrial invertebrates Diptera 5 ± 2 9 ± 1 19 ± 4 2 5 ± 1 Hemiptera 31 ± 9 59 ± 10 86 ± 13 49 ± 6 88 ± 9 Coleoptera 1 4 ± 1 2 3 ± 1 Hymenoptera 3 ± 1 9±2 10 ± 1 3 ± 1 3 ± 1 Lepidoptera + + + 7 ± 1 3 ± 1 Collembola 44 ± 22 39 ± 9 105 ± 26 71 ± 17 69 ± 15 Total 93 ± 35 123 ± 21 272 ± 50 137 ± 20 175 ± 23 Total, all groups 238 ± 73 324 ± 41 815 ± 101 1007 ± 95 537 ± 43 107 Johansen et al.: Diel fluctuations of invertebrate drift in a Norwegian stream significantly more during the day than at «night» A similar observation has been made in other stud (F(2,3wI4.25, P 40, 30 (a) I (c) c<)r---.. 30 1 T I 20 8 20 0 0 ..... I .I. J. J. 10 ~ 10 '" IJ) 0.. en 01 ~ 0 IJ) ..0 8 ::l 700 ~ 120] I (b) I (d) •c;j 90 525 t:: IJ) "'0 60 ~ I '\.I/I"-...T 1 T ~ 350 ~ 'q Cl 30 ~ 175 o+-I-~~---~~----~~--~ o +-1-~~-~-_~-~-_~-~-~~ 01 04 07 10 13 16 19 22 01 04 01 04 07 10 13 16 19 22 01 04 Time of day 3 Figure 3. Drift density (number of individuals per 100 m ) of Baetis spp. in Sooterelva. Notice the difference in scale between each month. Vertical bars represent 1 SE. Samples from (a) 22-24 May, (b) 23-25 June, (c) 18-20 August, and (d) 6-7 October 1996. Shaded bars on the x-axes represent period of darkness between sunset and sunrise. 108 Norw. J. Entomol. 47, 101-112.2000 range of values recorded for temperate and sub tic streams (Muller 1965, 1966). A lack of fluctu tropical streams (Waters 1972, Brittain & Eikeland ations in the drift is also found in fishless streams, 1988). In a study from Nordfjord, western 0-150 e.g. high altitude, tropical Andean (Turcotte & 3 individuals per 100 m • From Vindelalven in Swe Harper 1982, Flecker 1992) and Himalayan den, Ulfstrand (1968) found drift densities of200 streams (Brewin & Ormerod 1994). 3 500 per 100 m , similar to the range of96-663 per The diel drift pattern in October was a typical 100 m3 found in S 12 30 (a) (c) 9 20 10 24 (d) 18 12 6 01 04 07 10 13 16 19 22 01 04 01 04 07 10 13 16 19 22 01 04 Time of day 3 Figure 4. Drift density (number of individuals per 100 m ) of water mites (Hydracarina) in Sceterelva. Notice the difference in scale between each month. Vertical bars represent 1 SE. Samples from (a) 22 24 May, (b) 23-25 June, (c) 18-20 August, and (d) 6-7 October 1996. Shaded bars on the x-axes represent period of darkness between sunset and sunrise. 109 Johansen et al.: Diel fluctuations of invertebrate drift in a Norwegian stream depressed the drift. Small changes in light inten Emerging aquatic insects were most abundant in sity can affect drift, for example moonlight has drift samples in June and August, with adult Chi been shown to have a depressant effect on drift in ronomidae dominant. Most orders of aquatic in some rivers (Waters 1962, Anderson 1966, Bishop sects have their most intense emergence periods & Hynes 1969, Hynes 1975). This explanation is in these months (Brittain 1982, Pinder 1986, Lil however not in accordance with the results of lehammer 1988). On all sampling occasions, sig Elliott (1967) who found no evidence for the de nificantly more adult Chironomidae were taken pressant effect ofmoonlight on the drift in a small during the day than at night. Most workers have stream in southwest England. found highest drift density of emerging aquatic insects after sunset (e.g. Elliott 1967, Elliott & The die! fluctuations in the drift of aquatic inver Minshalll968, Dudgeon 1983). tebrates observed in August and October were eliminated in the continuous light ofthe polar sum Terrestrial drift has been shown to increase in mer. This was also reported by Muller (Muller sunny weather (Elliott 1967), intense rainfall, and 1965, 1966) and Hinterleitner-Anderson et al. flooding (O'Hop & Wallace 1983), so an increase (1992). Drift behaviour may be influenced directly in density ofdrifting terrestrial invertebrates could by selection, and the minimisation ofday drift may be expected to occur with the afternoon increase be adaptive (Kohler 1985). It has been suggested in discharge in May. The drift ofterrestrial inver that the evolution of a periodicity in drift was a tebrates was highest during the day on all sam predator avoidance mechanism to minimise mor pling occasions (although not significant). This tality due to fish predation (Allan 1978, 1984, agrees with the observations of other workers Flecker 1992) or invertebrate predation (Corkum (Elliott 1967, Turcotte & Harper 1982 & Pointing 1979). Studies in fishless streams have Terrestrial invertebrates contributed a significant found no evidence for consistent diel periodicities percentage to the drift in Sreterelva (7-39 %). This (Turcotte & Harper 1982, Flecker 1992, Brewin is similar to Elliott (1967) (9-38 %) and slightly & Ormerod 1994, Douglas et al. 1994). However, higher than in other studies (e.g. Bishop & Hynes experimental manipulations of fish populations 1969: 3 %; Hynes 1975: 1 %; Turcotte & Harper have shown no effects on drift periodicities (Allan 1982: 13-23 %; Waters & Crawford 1973: 8 %). 1982, Flecker & Allan 1984), apparently due to The high percentage ofterrestrial invertebrates is the large temporal scales over which adaptive re probably a result ofthe dense riparian vegetation sponses develop (Flecker 1992). As the threats along Sreterelva. Mason & McDonald (1982) from predation would be continuous in the Arctic found input ofterrestrial invertebrates to be depen summer, there are no optimal times for inverte dent upon the riparian vegetation, with higher in brates to be drifting and therefore no reason to put under alder trees than in the open. retain a periodicity. The lack ofperiodicity in the drift ofaquatic inver Most taxa followed the general pattern of being tebrates observed in the present study during the night-active in August and October but lacking summer exemplifies one ofthe problems with the any diel rhythm in May and June. The major ex drift classification of Waters (1972). As the low ception to this pattern was water mites (Hydra level daytime drift of night-active invertebrates carina), which were day-active on all sampling is termed «constant drift» and night-time drift is occasions. This has also been observed by other called «behavioural», it becomes impossible to workers in streams (Elliott 1967, Elliott & distinguish these two categories of drift in peri Minshalll968, Bishop & Hynes 1969, Waringer ods of constant light. This classification is useful 1992) and also lakes (Moon 1940, Pieczynski in recognising the multifaceted nature ofthe drift. 1964). A predator with a visual foraging behav However, it can also lead to the expectation that iour such as water mites may be influenced by the drift of all invertebrates conform to one of the need for light, thus resulting in an increased these categories. However, the establishment of daytime activity. further categories would not be particularly use 110 Narw. J. Entama!. 47, 101-112.2000 ful and would only lead to confusion (Brittain & of the invertebrate fauna in a stream ecosystem. Eikeland 1988). Arch. Hydrobiol. 66, 56-90. Brewin, P.A & Ormerod, S.l 1994. Macroinvertebrate In conclusion, this study gives support to the drift in streams ofthe Nepalese Himalaya. Freshwat. theory that predation risk probably is part of the BioI. 32, 573-583. reason behind diel fluctuations in drift of aquatic Brittain, lE. 1982. Biology of mayflies. Ann. Rev. Ent. invertebrates. There was a lack of fluctuations in 27,119-147. the midsummer period when light was continu Brittain, lE. & Eikeland, T.l 1988. Invertebrate drift ous, and a clear difference between night and day a review. Hydrobio1. 166,77-93. in August and October when nights were dark. Brekken, T., Fjellheim, A & Larsen, R. 1981. Drift of benthic animals into and out of the weir basin at This suggests that the invertebrates use the dark Ekse. - The Weir Project, Information Bulletin, ness at night as a refugium from drift-feeding fish. Norwegian Water Resources and Electricity Board 14,1-37. (In Norwegian with English summary) Acknowledgements. We are most grateful for the help Clifford, H.F. 1972. A year's study of the drifting or of Henning Syvertsen, Torfinn Sorensen and Thomas ganisms in a brown-water stream of Alberta, 01sen in the field. We would also like to thank John O. Canada. Can. J. Zool. 50, 975-983. Solem and an anonymous referee for valuable com Corkum, L.D. & Pointing, PJ. 1979. 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Sandlund, O.T 1982. The drift of zooplankton and Hynes,1.D. 1975. Downstream drift of invertebrates microzoobenthos in the river Strandaelva, western in a river in southern Ghana. Freshwat. BioI. 5, 515 Norway. Hydrobiol. 94, 33-48. 532. Schreiber, E.S.G. 1995. Long-term patterns ofinverte Kohler, S.L. 1985. Identification ofstream drift mecha brate stream drift in an Australian temperate stream. nisms, an experimental and observational approach. Freshwat. BioI. 33, 13-25. Ecology 66, 1749-1761. Statzner, B., Dejoux, C & Elouard, 1.-M. 1984. Field Lillehammer, A. 1988. Stoneflies (Plecoptera) ofFenno experiments on the relationship between drift and scandia and Denmark. Fauna Ent. Scand. 21, 1-165. benthic densities ofaquatic insects in tropical streams Lillehammer, L., Raastad, l.E., Pethon, P., Barstad, G., (Ivory Coast). I. Introduction: a review of drift lit Lunde, V. & Onstad, S. 1995. Studier av bunn- og erature. methods, and experimental conditions. 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Developments in stream ecosystem Ulfstrand, S. 1968. Benthic animal communities in theory. Can. 1. Fish. Aquat. Sci. 42, 1045-1055. Lapland streams. Oikos, Suppl. 10, 1-120. Moon, P.P. 1940. An investigation of the movements Vannote, R.L., Minshall, G.w., Cummins, K.W., Sedell, of freshwater invertebrate faunas. - 1. Anim. Ecol. 1.R. & Cushing, CE. 1980. The river continuum 9,76-83. concept. Can. l. Fish. Aquat. Sci. 37,130-137. Mu1adal, H. & Skotvold, T 1993. Undersokelse av Waringer, J.A. 1992. The drifting of invertebrates and forurensningsforhold i Mokelandsvassdraget. 325/ particulate organic matter in an Austrian mountain 5/93, 1-22. (In Norwegian) brook. Freshwat. BioI. 27, 367-378. Mii1ler, K. 1965. Field experiments on periodicity of Waters, TF. 1962. Diurnal periodicity in the drift of freshwater invertebrates. Pp. 314-317 in Aschoff, stream invertebrates. Ecology 43, 316-320. 1. (ed.) Circadian clocks. North-Holland Publish Waters, TF. 1965. Interpretation of invertebrate drift ing, Amsterdam. in streams. Ecology 46,327-334. Muller, K. 1966. Die Tagesperiodik von Fliesswasser Waters, TF. 1969. Invertebrate drift - ecology and sig organismen. Z. Morph. Okol. Tiere 56, 93-142. nificance to stream fishes. Pp. 121-134 in Northcote, Muller, K. 1970. Tages- und Jahresperiodik der Drift T.G. (ed.) Symposium on salmon and trout in in Fliessgewassern in verschiedenen geographi streams. Univ.British Columbia, Vancouver. schen Breiten. Oikos Suppl. 13, 21-44. Waters, TF. 1972. The drift ofstream insects. Ann. Rev. Muller, K. 1974. Stream drift as a chronobiologica1 Ent. 17, 253-272. phenomenon in running water ecosystems. Ann. Waters, TF. & Crawford, G.w. 1973. Annual produc Rev. Ecol. Syst. 5, 309-323. tion of a stream mayfly population: a comparison O'Hop,1. & Wallace, 1.B. 1983. Invertebrate drift, dis of methods. Limnol. Oceanogr. 18,286-296. charge, and sediment relations in a southern Appa lachian headwater stream. Hydrobiol. 98, 71-84. Received 8 January 2000. Pearson, W.D. & Franklin, D.R. 1968. Some factors accepted 10 May 2000 112 Norw. J. Entomol. 47, 113-124. 2000 Life cycle, diet and habitat of Polycentropus f1avomaculatus, Plectrocnemia conspersa and Rhyacophila nubila (Trichoptera) in I2Jvre Heimdalen, Jotunheimen Mountains, Norway Sigve Reiso &John E. Brittain Reiso, R. & Brittain, lE. 2000. Life cycle, diet and habitat of Polvcentropus flavomaculatus, Plectrocnemia conspersa and Rhyacophila nubila (Trichoptera) in 0vre Heimdalen, Jotunheimen Mountains, Norway. Norw. 1 Entomo!. 47,113-124. Life cycles, diets and habitats ofthe three trichopterans, Polycentropusflavomaculatus (Pictet, 1834), Plectrocnemia conspersa (Curtis, 1834) (Polycentropodidae) and Rhyachopila nubila (Zetterstedt, 1840) (Rhyachophilidae) were investigated. All three species showed indications of semivoltine life cycles, probably due to low temperatures. P flavomaclllatus dominated the outlet of the lake, 0vre Heimdalsvatn, while R. nubila was widely distributed, but dominated the faster flowing reaches. P cOllspersa dominated in the pools in the upper reaches of the stream Brurskardbekken. Both P flavomaclllatus and P conspersa were common in the outlet of the small lake Brurskardtjem. The dietary composition of the three predaceous trichopteran species reflected a combination of prey occurence and foraging/capture techniques. The free-living R. nubila had the narrowest niche breadth while the net-spinning, P conspersa and P flavomaculatus had wider niche breadths. How ever, the species had several similar prey taxa in their diets, indicating that overlap also occurred between predators of very different foraging/capture modes. They all showed monthly variations in feeding habits as a result of seasonal variations in the abundance of prey, indicating that these spe cies are generalists. This coupled with their ability to change to semivoltine life cycles at high altitudes is probably an important factor explaining their wide geographical distribution. Keywords: Trichoptera, life cycle, habitat, diet. Sigve Reiso, Department ofBiology and Nature Conservation. Agricultural University ofNonvay. N-1432 As, Nonvay. [Present adress: Ragna Nielsensvei 12. N-0592 Oslo. Norwa~j. John E. Brittain, Zoological Museum, University olOslo. Sars gate 1, N-0562 Oslo, Norway. INTRODUCTION distributed representatives of two primarily pre daceous families, the Polycentropodidae and the Certain trichopteran species demonstrate flexible Rhyacophilidae, respectively (Dudgeon & Ri life cycles throughout their distributional range, chardson 1988, Wiggins 1996). Pflavomaculatus shifting from a univoltine life cycle in the low and P conspersa (Polycentropodidae) catch their lands to a semivoltine cycle in montane areas prey from the drift using elongate, sack-like nets (Solem & Gullefors 1995). Towards their altitudi (Wiggins 1996). Both species are found through nallimits, semivoltine life cycle strategies are more out Europe (Malicky 1979, Szczesny 1986, Solem likely in aquatic insects, mainly due to low water & Gullefors 1995). R. nubila (Rhyacophilidae) is temperatures (Brittain 1978, 1983, 1990, Ward & free living and actively searches for its prey. It Stanford 1982, Sweeney 1984). has a northern distribution in Europe and Russia Polycentropus flavomaculatus, Plectrocnemia (Fjellheim1976, Solem & Gullefors 1995). conspersa and Rhyacophila nubila are three widely In this present study, life cycle, habitat and diet of 113 Reisa & Brittain: Life cycle, diet and habitat af Trichaptera these three widely distributed trichopteran species bekken, and one station in the outlet of the lake, were investigated in a mountainous area. The aim 0vre Heimdalsvatn, were studied. Brurskard ofthis study was to obtain knowledge oftheir ecol bekken rises at an elevation of 1309 m a.s.\. in a ogy at or near their altitudinallimits. small lake, Brurskardtjern, located at the upper limit ofthe low alpine zone. Subalpine vegetation SITE DESCRIPTION and mountain birch (Betula pubescens) reach alti tudes of about 1200 m a.s.\. along Brurskard This study was carried out in 0vre Heimdalen, a bekken. It falls gradually until 1275 m a.s.\., where valley on the eastern slopes of the Jotunheimen the stream flows slowly through a series of small Mountains in southern Norway (Latitude 61 °25'32" pools. After 1250 m a.s.\. the gradient increases N) (Figure I). The valley and the lake 0vre Heim substantially until the stream reaches the val1ey dalsvatn ecosystem were thorougly described in Vik floor at 1100 m a.s.\. The substrate ofthe streambed (1978). Four stations in the stream, Brurskard in the upper parts of the river is mostly bedrock ~w-?"'n.t'/r"', t;J t- 60'~ V SOOm 1000m. lS00m Figure 1. Map of 0vre Heimdalsvatn and the stream Brurskardbekken. Stations 1 to 5 are marked. Altitudes in m a.s.\. 114 Norw. J. Entomol. 47, 113-124.2000 and large stones while smaller stones and sand Brurskardbekken (station 3 and 4) and one at the dominate the slow flowing lower parts, From the outlet of0vre Heimdalsvatn (station 5). An ultra Brurskardtjern, the stream flows for 3.2 km be violet light trap was used on four occasions at sta fore entering the lake, 0vre Heimdalsvatn, at 1090 tion 5. Adults were also collected in sweep-net m a.s.l. The outlet of 0vre Heimdalsvatn is 5-10 samples ofriparian vegetation at all stations. Adult m wide and 1.5 m deep. The water is slow flow collections were used to establish emergence and ing with some large pools and the substrate is com flight periods. posed mainly of large stones. The location of the All samples were preserved in 70 % alcohol in study reaches is given in Figure 1. More details the field, then sorted and identified in the labora concerning the reaches are given in Reiso (1999). tory to species using a dissecting microscope. The Water temperatures were recorded by digital log larvae were identified to genus using Solem & Gul gers in Brurskardbekken (station 4) and the outlet lefors (1995) and to species from Edington & Hild of0vre Heimdalsvatn (station 5) every hour, and rew (1981). Adults were identified using Malicky every 3 hours respectively, during the ice-free pe (1983). For life history analysis, larval head cap riod (Figure 2). The mean summer temperature sule length and width was measured using an ocu (June-September) was 9.6 QC at station 5 and 7.1 lar micrometer at 40x magnification. QC at station 4. Where possible the foregut offive, fifth instar lar vae from each of the three species was examined METHODS on six sampling occasions (Figure 5). The fore Larvae, prepupae and pupae were collected at all guts were removed and mounted on glass slides. five stations on seven occasions during the ice After mounting in Hoyers Medium the relative free period from June to October 1998. They were contribution ofdifferent food items was estimated collected by a kicking technique (Frost et al. 1971) using an ocular squared grid. The percentage area (30x30 cm net, mesh size 350 !-tm) and hand picked occupied by each prey category was calculated. from stones and other substrates in order to col Eleven categories were used: Copepoda, Clado lect all stages. Three, 30 sec. kick samples were cera, Hydracarina, Chironomidae, Simuliidae. taken at each station for density estimates. Ephemeroptera, Plecoptera, Trichoptera, Coleo ptera, adult insects and detritus. Guts with no con Adults were collected by a variety of methods. tents were classified as empty. Three different malaise traps was used. two by \ ,', ..' :. I ...... -StS Sl41 I 'uA A ' ,..... ' '.,,.:.' .. ', .. \' , \ , ..'.:': , ,.' I /~,.:'I· ".4" ,:~. • '\,.,\:\ ....,., .'1 t' It~, ~,:\ \;~ J/!"0 I" ... • .' I ), • '0' '" _ .. ; .'~ ,'...: ~ ~\' \ .... \ . ,. 4" '-,, . 4 .- ,.' .. ",.!""':' 2 , ,":,. ~"..; .". o 21/5 416 18/6 217 1617 3017 13'8 27/8 10'9 24/9 Figure 2. Mean daily water temperatures in Brurskardbekken (station 4) and at the outlet of 0vre Heimdalsvatn (station 5) from 21 May to 3 October 1998. 115 Reiso & Brittain: Life cycle, diet and habitat of Trichoptera Niche breadth and niche overlap with respect to the formula: diet were calculated for each trichopteran species. n _I n 2 2 Oxy = CIPx.j Py,J)/(v LPx,J Pv,J) Niche breath (B) is given as: jd j=1 B=I/V/ where Px,J and P;;i are the percentage use by spe J = 1 cies x and y ofthe same prey group. Values> 0.75 where P j = proportion of the prey group j in the are indicative of high overlap between species total diet, and B can range from 0 - n (n = number (Dudgeon & Richardson 1988). of prey groups) (Dudgeon & Richardson 1988). Niche overlap (Oxy) was calculated according to P. flavomaculatus 2, n=421 E E 1,5 .!: .<:: ..,;I!"I rnc: .91 .,... "0 ~ ..,., 0,5 .... 01 E • , , , ° 0,5 1,5 2 2,5 P. conspersa 3,5 n=84 3 E E 2,5 ,..; .!: • .<:: 2 m ~ 1,5 .... • • "0 ca 1 Q) • :r: 0,5 ° 0,5 1 1,5 2 2,5 ° R. nubila 1,5 n = 106 E ~.I E • •••• .S ...... <:: m c: "",. Q) ~ 0,5 .. Figure 3. Head width and Q) _. -. :r: head length measured for • the different instars of P f1a vomaculatus, P conspersa 0,5 1,5 2 2,5 °0 and R. nubila from 0vre Head width in mm. Heimdalen during 1998. 116 Norw. J. Entomol. 47, 113-124.2000 RESULTS Head capsule measurements of larval P jlavo maculatus fell into discrete groups, representing the Polycentropus flavomaculatus five larval instars (Figure 3). The size distribution In 0vre Heimdalen, Pjlavomaculatus was found offifth instar larvae was bimodal, representing male at stations 1 and 5. The species constituted a large and female larvae. First instar larvae were found at part of the trichopteran fauna in these outlet areas the beginning of June and in August, but only in (Table 1). The P jlavomaculatus population in the low numbers (Figure 4). Second instar larvae were outlet of 0vre Heimdalsvatn had the highest den found in June and August, while third instar larvae sity ofTrichoptera at all stations, with an average occurred in June and July. Fourth and fifth instar of 19.9 larvae per kick sample during the ice free larvae were found throughout the ice-free period. period. At station 1 the average was 0.9 P jlavo Pupae were recorded throughout July, and adults maculatus larvae per 30 sec. kick sample. were taken from late July to mid-August. P. flavomaculatus 57 40 A ,. ,. p ~ pp 0 0 5 • 1 I 4 •• • • I •I 3 • •I • 2 1 •I i 26 34 30 40 118 129 44 P. conspersa A p pp5 t! ---._r---.iir-----.__ - - 4 3 2 1 13 10 16 35 R. nubila 11 52 17 Figure 4. Instar frequency distribu A f f f f t tion on each sampling date during p 0 <:::I 0 0 1998 for P. f1avomaculatus, P. con pp 0 ""0 "" spersa and R. nubi/a from 0vre 5 "" 11 Heimdalen. Numbers above dates 4 • •I • 11 • • are number of larvae used in the 3 • • 2 analysis. The width of each bar is 1 • •I proportional to the frequency of that 8 5 26 4 7 22 34 instar on that date. Numbers above 4 Jun. 16 Jun. 7 Jul. 21 Jul. 11 Aug. 12 Sept. 3 Oct. arrows are total number of adults collected. 117 Reiso & Brittain: Ufe cycle, diet and habitat of Trichoptera Table 1. The total number and the frequency composition (%) of the total trichopteran larval fauna for P flavomaculatus, P conspersa and R. nubila at each station in 0vre Heimdalen. ,. Station 1 2 3 4 5 P flavomaculatus 18(39.1) 418 (92.7) P conspersa 19(41.3) 36 (85.7) 2 (6.7) R. nubila 8 (17.4) 6 (14.3) 36 (72) 7 (53.8) 32 (7.1) The diet of P. jlavomaculatus was characterised Rhyacophila nubila by seasonal variation in the type offood consumed Rhyacophila nubila was found at all five stations. (Figure 5), having a niche breadth value of 6.65 It dominated the trichopteran fauna in the faster with respect to diet. Most groups ofprey were frag flowing inlet area at stations 3 (72 %) and 4 (53.8 mented when found in the foregut, although small %). At station 5, relatively high numbers of R. Chironomidae were sometimes found whole. P. nubila occurred, although only comprising 7.1 % jlavomaculatus had dietary overlap of 82 % and of the total trichopteran fauna (Table 1). 50 % withP conspersa and R. nubila, respectively. Head capsule measurement of larval R. nubila Plectrocnemia conspersa collected fell into five groups, representing dis crete larval instars (Figure 3). In contrast to the Plectrocnemia conspersa was found at stations 1, other two species there was no evidence ofsexual 2 and 3. At stations 1 and 2, it was the dominant dimorphism in the fifth instar. Fifth instar larvae species comprising 41.3% and 85.7% of the and pupae occurred on all sampling occasions trichopteran fauna collected in kick samples (Ta throughout the ice-free period. Only one first instar ble 1). At station 3 only two larvae were recorded larva was recorded, collected in August. Adults during the ice-free period. Head capsule measure were taken from beginning ofJuly to October, with ments of the larval P conspersa fell into three a peak during August (Figure 4). groups representing larval instars (Figure 3). No first or second instar larvae were recorded. One, The foregut of R.nubila larvae contained six dif third instar larvae was found in late July. Fourth ferent prey categories (Figure 5). All prey groups instar larvae were found from late July to Octo were highly fragmented. R. nubila had a niche ber. Fifth instar larvae were found on all sampling breadth value of2.37 and dietary overlap of 50 % occasions and showed a clear separation in size and 47 % with P jlavomaculatus and P conspersa, between male and female larvae. Pupae were re respectively. corded in July and adults in late July and August (Figure 4). DISCUSSION The diet of P conspersa was also characterised Life cycle by seasonal variations in the type of food consu Trichopteran species may have multivoltine, uni med (Figure 5). had a niche breadth P conspersa voltine or semivoltine life history strategies. How of 3.28 with respect to diet. The prey found in the ever, most species in Northwest Europe are either foreguts were less fragmented than in P jlavoma univoltine or semivoltine, while some are able to culatus and R. nubila. P. conspersa had dietary alternate between the two strategies depending on overlap of 82 % and 47 % with P .flavomaculatus the environmental conditions (Solem & Gullefors and R. nubila, respectively. 1995). At high altitudes or towards their northern limits, many widely distributed trichopteran spe cies require more than one year to complete their 118 Norw. J. Entomol. 47,113-124.2000 P. conspersa n=(Empty guts) 2 3 5(2) 5 5 5 ~ Hydracarnia • Terrestrial prey ClI e ~Cladocera ClI -o IIl!I Plecoptera 'E 8... mOetritus III Q. o Chironomidae • Simuliidae o% +-'----'- P. flavomaculatus n=(Empty guts) 5(1) 5(2) 5(2) 5(4) 5 5 o Copepoda 100 % 1 []Coleoptera • Terrestrial prey ClI 80% f ~Cladocera ClI 60% Ephemeroptera .c> a c: mPlecoptera - 40 % ~ mOetritus III Q. 20% o Chironomidae • Simuliidae o% +-'----'.~ R. nubila n= (Empty guts) 2 5 5 5 5 100 % .. • Trichoptera ClI 80 % 1 III... ~ Ephemeroptera ClI -0 III Plecoptera c: -III 40 %-1 mOetritus ...u Gl o Chironomidae Q. 20 % 1 .Simuliidae 0% 4 Jun. 7 Jul 21 Jut. 11 Aug. 11 Sept. 3 Qct. Date Figure 5. Monthly variations in dietary compositions of P f1avomaculatus, P conspersa and R. nubila in 0vre Heimdalen during 1998. The percentage area occupied by each prey category in fore-guts mounted on glass slides, is displayed. Numbers above bars are total guts analysed on that date. 119 Reisa & Britlain: Life cycle, diet and habitat af Trichaptera life cycle (Andersen & Tysse 1984). This seems (Hildrew & Wagner 1992), while in another study to be the situation for P. jlavomaculatus, P con pupae metamorphosis varied between 12 days at spersa, and R. Ilubila in 0vre Heimdalen. 16-17 °C to 23 days at 10-11 °C (Tachet 1967). At station 2, where P. conspersa was found in All instars of P. jlavomaculatus were present at highest densities, Sand (1997) recorded the mean the beginning of June, indicating that the larvae temperature to be 8.9 °C during the ice-free pe of this species overwinter in all stages (Figure 4). riod. Considering the low summer temperatures This fact, together with the presence offifth instar in Brurskardbekken, a semivoltine life cycle seems larvae in all samples during the ice-free period most likely. and a restricted emergence period, suggests a semivoltine life cycle. In an earlier study, Eikeland The life cycle ofR. nubila is more complex, with (1982) found that P jlavomaculatus population fifth instar larvae present during summer and a in 0vre Heimdalen varied between a two and a long emergence period (Figure 4). The presence three year life cycle. He mainly explained this offirst and second instar in August indicates hatch semivoltine life cycle by the low temperatures. In ing of a new generation. Fjellheim (1976) found another low temperature location in northern Swe the egg incubation time for R. Ilubila to be 16 days den, Ulfstrand ( 1968) also considered the species at 18°C. However, Brurskardbekken temperatures to have a two year life cycle. In contrast, Recasens were around 10°C at the beginning ofAugust and & Puig (1987) found P. jlavomaculatus to have the incubation period would be longer. The presen three generations per year in northern Spain, while ce ofadults in September indicated oviposition at Elliott (1968) found P. jlavomaculatus to be uni even lower temperatures leading to overwintering voltine in southern England. In close association in early instars or even as eggs. These overwinte with the influence ofwater temperature, the length ring early instar larvae or eggs probably needed of the ice-free season and the abundance of prey an additional winter before emergence, leading also influence the variation in life cycle length. to a semivoltine life cycle. In Swedish Lappland, Ulfstrand (1968) considered R. nubila to have a P. jlavomaculatus emerged during late July and semivoltine life cycle, while in western Norway, early August. The presence of first and second Fjellheim (1976) found R. nubila to be univoltine. instar larvae in August suggested that egg incu Both studies found life cycles difficult to inter bation had a duration of about one month. During pret due to poor synchronisation. this period water temperatures were around 12°C in the outlet of 0vre Heimdalsvatn. This fits in The presence ofR. Ilubila pupae early in the sea well with the laboratory results ofEikeland (1982) son indicated that some larvae overwinter as pu in which a mean incubation time of34 days for P. pae. This phenomenon has also been found in the jlavomaculatus eggs at 12°C was found. related species Rhyacophila dorsalis (Edington & Hildrew 1981). The pupae would have continued The life history ofP. compersa is not readily ap to develop when water temperatures rose, emerg parent because too few larvae were collected. ing in early summer. This might be one ofthe rea However, the available data also suggest a semi sons why R. nubila had such a long flight period, voltine life cycle for this species, due to the pres lasting from early July until October with a peak ence of fifth instar larvae on all sampling occa in early August. A long flight period has also been sions and a restricted emergence period thus re shown for R. nubila in the Dovrefjell Mountains sembling the life cycle of Pflavomaculatus (Fig (Solem 1985) and in western Norway (Fjellheim ure 4). Szczesny (1986) and Lancaster et al. (1995) 1976). assumed P. conspersa to be univoltine in Poland and England, respectively. Water temperatures and food quality have been considered the most important factors prolong Studies ofegg and pupal development have shown ing life cycle in aquatic insects (Brittain 1978, how temperature affected the life cycle length of 1990, Ward & Stanford 1982, Sweeney 1984). Re P. conspersa. The egg development period varied viewing the benthic insects in the lake 0vre Heim between 93 days at 6 °C and 20 days at 18°C 120 Norw. J. Entomol. 47,113-124.2000 dalsvatn, Larsson et al. (1978) found only one ofstreams. The upper reaches of a stream may be example of a multivoltine life cycle. In contrast, more favourable for a larger species like P con semivoltine life cycles were fairly common in this spersa because in these areas there are more larger high altitude area. Andersen & Tysse (1984) found potential prey items compared to the lake outlets. that the limnephilid, Chaeopteryx villosa, had a Otto (1984) found the dominant size group ofdrift univoltine life cycle in a lowland stream (90 m ing potential prey animals to be three times larger a.s.!.) and a semivoltine life cycle in a mountain in a stream than in a lake outlet, suggesting a stream (1220m a.s.l.) in western Norway. Brittain mechanism whereby prey size might be a factor (1978) described a similar pattern for the stonefly affecting their distribution. Nemurella pictetii. Such adaptations enable the Temperature may be a factor influencing distri species to colonise a wider range of habitats than bution patterns of polycentropodid larvae. P jla species with fixed life cycles. Species with a flex vomaculatus was only found in high numbers in ible life cycle will also be able to alternate life cy lake outlets, areas having a relatively higher mean cles and survive in unstable and harsh environments temperature than streams (Richardson and Mac such as mountain areas. The semivoltine life cycle kay 1991). However, at station 1 both species oc strategies of P jlavomaculatus, P conspersa and curred together, showing that P jlavomaculatus R. nubila are therefore an adaptation to the rela can live under the same temperature regime as P tively low summer temperatures and the eight conspersa and that temperature might not be the months ofice and snow cover in 0vre Heimdalen. most important factor governing the distribution Few first instar larvae were found during this of the two species in 0vre Heimdalen. study. This might be due to short duration and The habitat ofR. nubila was different than for the rapid development in the early instars during the two polycentropodid species. It occurred at all the summer or different habitat use between instars. sampling stations in 0vre Heimdalen, but was In western Norway, Fjellheim (1976) found that most numerous at stations 3 and 5, station 3 be R. nubila developed rapidly from egg to second ing a typical fast flowing area while station 5 was instar in the autumn, before ice cover. The same slow flowing. This indicates little preference for rapid development from egg to second instar was water velocity in R. nubila as found in the two also found in the P jlavomaculatus population in polycentropodid species. However, B~kken et al. 0vre Heimdalen (Eikeland 1982). (1984) demonstrated that R. nubila disappeared from a former riffle after the construction ofa weir Habitats dam, suggesting the species has a water velocity Pjlavomaculatus, P conspersa andR. nubila oc threshold. In high velocity areas like stations 3 cupied different habitats in the streams of 0vre and 4, R. nubila was the dominant Trichoptera spe Heimdalen (Table 1). P jlavomaculatus and P cies. Its ability to exploit such habitats reduces conspersa were typical pool species, inhabiting competition with other predaceous trichopterans areas oflow current velocity (Edington 1968). P like P jlavomaculatus and P conspersa in 0vre jlavomaculatus was common in the two lake out Heimdalen. lets (stations 1, 5), while P conspersa was com mon in the lake outlet and pools in the upper Diet reaches of Brurskardbekken (stations 1,2). Ear The diet ofthe three predaceous trichopteran spe lier studies have shown that P conspersa usually cies (Figure 5) reflected a combination ofthe prey inhabits tributaries while P jlavomaculatus oc found in the habitat and the foraging/capture tech curs in the main stream (Otto 1985). Otto (1985) niques used. P jlavomaculatus and P conspersa suggested that P conspersa dominates over P are typical filter feeders, passively catching prey jlavoma~ulatus when their distributions overlap drifting/crawling into their nets. By contrast R. because of its larger size. This might be the rea nubila actively searches for its prey, lacking con son why P conspersa dominates the upper reaches 121 Reiso & Brittain: Life cycle, diet and habitat of Trichoptera structions to aid prey capture. However, the over coptera and Trichoptera hatched from eggs laid all similarity ofthe diets ofthe study species was earlier in the summer and early instars of these notable. All three species consumed Simuliidae, two groups seem to be ofmajor importance in the Chironomidae and Plecoptera although there was R. nubila diet. The presence ofdetritus in late sum interspecific variation in the importance of these mer could also be a sign of lower prey densities items. The diets of the two closely related P. and reduced activity ofboth R. nubila and its prey jlavomaculatus and P. conspersa were most simi due to lower water temperatures. lar (overlap 82 %). However, R. nubila exhibited P. jlavomaculatus and P. conspersa consumed considerable overlap in diet with the two poly similar types of prey including both aquatic and centropodid species indicating their overlap also terrestrial items. The large difference in niche occurred between predators ofvery different fora breath between the two species was probably asso ging/capture mode. Townsend & Hildrew (1979) ciated with a lack of dietary dominance by a sin found considerable overlap between P. consper gle prey group in the diet ofP. jlavomaculatus. In sa and the more mobile alderfly Sialisfuliginosa, the diet of P. conspersa, Cladocera were clearly indicating the same pattern. On the basis of high the most important prey item during August, Sep dietary overlap in the polycentropodid species, it tember and October. The dominance ofPIecoptera is possible that competition for food and for suit in the guts of P. jlavomaculatus in June could be able net spinning sites could be a factor influenc due to the life cycle of this species. Many pleco ing their local distribution. pteran species emerge during June (Lillehammer The monthly variations in feeding habits of the 1978) and therefore have their maximum activity larvae (Figure 5) reflected the seasonal variations in this month. The presence of Copepoda in the in prey abundance. For example, in early July, all June sample can be explained by the flushing of three trichopteran species showed a dominance the plankton population in the lake during the ofsimuliid prey in their diet, indicating a simuliid spring snowmelt spate (Larsson 1978). peak in the benthos at that time. This fits with Chironomids occurred in guts of both P. flavo observations of dense simuliid aggregation on maculatus and P. cOllspersa in June, August and rocks in all parts ofthe stream during July. Bildeng September. This fitted emergence pattern ofchiro (1982) found the same pattern in his study of the nomids along the shores of 0vre Heimdalsvatn, filter feeding trichopteran, Arctopsyche ladogen where they showed peak emergence in June and sis, in the River Glomma. In June 50 % of the August (Brittain & Lillehammer 1978). The pres benthic fauna by numbers consisted of simuliid ence ofCIadocera in the gut ofP.jlavomaculatus larvae, leading to a dominance of simuliid in the in July was due to the life cycle of Holopedium species gut. gibberum. This species hatched from resting eggs R. nubila had the narrowest niche breadth with at the beginning of July and reached maximum respect to diet, consuming six prey categories. The densities during July in 0vre Heimdalsvatn (Lars lack of zooplankton in the diet reflects its forag son 1978). ing/capture techniques. It detects its prey by vi Wind and rain can explain the occurrence of adult sion or sound, while it explores the substrate us insects in the diet in late July. Such weather condi ing side to side motions (Malmqvist & Sackmann tions may cause flying and emerging insects to fall 1996). During June and July, the diet was domi down into the water where they can be captured in nated by Chironomidae and Simuliidae, both nu Pjlavomaculatus and P. conspersa nets. Townsend merous in Brurskardbekken (Lillehammer & Brit & Hildrew (1979) found terrestrial prey in P COIl tain 1978). After July other prey categories be spersa guts in significantly greater numbers after come important. This can be explained by the a period of high winds and rain. In late July and in emergence of Simuliidae and Chironomidae in August, adult insects were more important for P. mid summer, forcing R. nubila to change to other conspersa than for P. jlavomaculatus. This could prey groups. In late summer early instars of Ple be due to greater wind exposure in the area around 122 Norw. J. Entomol. 47, 113-124.2000 stations 1and 2 compared to the main valley where animal production in a weir basin area in western the outlet is situated. Norway. Pp. 223-232 in Lillehammer A. & Saltveit S. 1. (Eds), Regulated rivers. University Press, Oslo. In late summer Bosmina longispina, reached its Dudgeon, D. & Richardson, 1. S. 1988. Dietary varia maximum densities in 0vre Heimdalsvatn (Lars tions of predaceous caddisfly larvae (Trichoptera: son 1978) and Cladocera were found in the diet Rhyacophilidae, Polycentropodidae and Arctopsy ofboth polycentropodid species. The relative high chidae) from British Columbian streams. Hydro amount ofempty guts found in P. jlavomaculatus bio!. 160,33-43. in July and August could be due to fifth instar Edington, J. M. 1968. Habitat preferences in net-spin larvae preparing for pupation. A similar pattern ning caddis larvae with special reference to the in fluence of water velocity. 1. Anim. Eco!. 37, 675 was found in P. conspersa. 692. In conclusion all three species showed monthly Edington, J. M. & Hildrew, A. G. 1981. A key to to the variations in feeding habi ts as a result ofseasonal caseless caddis larvae ofthe British Isles, with notes variations in the abundance of the prey, indicat on their ecology. Sci. Pubis. Freshw. Bio!. Ass. 43, 1-92. ing the species to be generalists. Eikeland, T J. 1982. Livssyklus og fodeopptak hos Their flexible life cycle and the fact that they are Polycentropus flavomaculatus (Pictet) (Trichopte generalists with regard to diet are probably im ra) i 0vre Heimdalsvatn, 0st-Jotunheimen (In Nor portant factors explaining their wide geographi wegian). 64 pp.Thesis, Zoological Museum, Uni versity of Oslo, Norway. cal distribution. Elliott, J. M. 1968. The life histories and drifting of Trichoptera in a Dartmoor stream. 1. Anim. Eco\. Acknowledgements. We wish to thank Gunnar Hal 37,169-182. vorsen for valuable comments on the manuscript, and Fjellheim, A. 1976. Livssyklus, produksjon og drift hos John Solem for important information on some of the Rhyacophila Ilubila (Zett.) (In Norwegian). 106 pp. species. Research facilities in 0vre Heimdalen were Thesis, Zoological Museum, University ofBergen, provided by the Zoological Museum, University of Norway. Oslo. The Norwegian Water Resources & Energy Di Frost, S., Huni, A. & Kershaw, W. E. 1971. Evaluation rectorate (NVE) kindly supplied temperature data from of a kicking technique for sampling stream bottom the outflow of 0vre Heimdalsvatn. fauna. Can. 1. Zoo!. 49, 167-173. Hildrew, A. G. & Wagner, R. 1992. The briefly colo References niallife ofhatchlings of the net-spinning caddisfly Plectrocllemia conspersa. J. N. Am. Benth. Soc. Andersen, T & Tysse, A. 1984. Life cycle of Chaeto I L 60-68. pteryx villosa (Fabricius, 1798) (Trichoptera: Lim Lancaster, J. & Robertson, A. L. 1995. Microcrustacian nephilidae) in a lowland- and a Mountain- Stream prey and macroinvertebrate predators in a stream in Western Norway. Aquatic Insects 6, 217-232. food web. Freshw. Bio!. 34,123-134. Bildeng, R. 1982. Livssyklus og nreringsanalyse hos Larsson, P. 1978. The life cycle dynamics and produc Arctopsyche ladogensis (Kolenati) (Trichoptera) i tion ofzooplankton in 0vre Heimdalsvatn. Holarc. Glomma (In Norwegian). 77 pp. Thesis, Univer Eco!. I, 162-218. sity of Oslo, Norway. Larsson, P., Brittain, J.E., Lien, L., Lillehammer, A. & Brittain, J. E. 1978. Semivoltism in mountain popula Tangen, K. 1978. The lake ecosystem of 0vre tioms ofNemurella pictetii (Plecoptera). Oikos. 30, Heimdalsvatn. Holarc. Eco!. 1,304-320. 1-6. Lillehammer, A. 1978. The Plecoptera of 0vre Heim Brittain, J. E. 1983. The influence of temperature on dalsvatn. Holarc. Eco!. 1, 232-238. nymphal growth rates in mountain stoneflies (Ple Lillehammer, A. & Brittain, J. E. 1978. The inverte coptera). Ecology 64,440-446. brate fauna of the streams in 0vre Heimdalen. Brittain, J. E. 1990. Life history strategies in Epheme Holarc. Eco!. I, 271-274. roptera and Plecoptera. Pp. 1-12 in I. C. Cambell Malicky, H. 1979. Notes on some caddisfly (Trichopte (ed.), Mayflies and Stoneflies: Life Histories and ra) from Europa and Iran. Aquat. Insects 1, 3-16. Biology. Kluwer Academic Publishers, Dordrecht, Malicky, H. 1983. Atlas of European Trichoptera. 298 Netherlands. pp. Dr W. Junk Publishers, The Hauge-Boston-Lon Brekken, T, r)ellheim, A. & Larsen, R. 1984. Benthic don. 123 Reiso & Brittain: Life cycle, diet and habitat of Trichoptera Malmqvist, B. & Sackmann, G. 1996. Changing risk Rev. Entomol. 27, 97-117. ofpredation for a filter-feeding insect along a cur Wiggins, G. B. 1996. Larvae of the North American rent velocity gradient. Oecologica l08, 450-458. Caddisfly Genera (Trichoptera), 2 nd. ed. 457 pp. Otto, C. 1985. Prey size and predation as factor gover University of Toronto Press, Toronto. ning the distribution of lotic polycentropodid cad disfly larvae. Oikos 44, 439-447. Received 16 June 2000, Otto, C. 1993. Long-term risk sensitive foraging in accepted 31 October 2000 Rhyacophila nubila (Trichoptera) larvae from two streams. Oikos 68, 67-74. Recasens, L. & Puig, M., A. 1987. Life cycles and growth patterns of Trichoptera in the Matarrana, a karstic river. Series Ent. 39,247-251. Reiso, S. 1999. Life cycle, diet and habitat of Polycen tropus flavomaculatus, Plectrocnemia conspersa and Rhyacophila nubila (Trichoptera) in 0vre Heimdalen of the Norwegian Jotunheimen Moun tains. 2l pp. Thesis, Agricultural University ofNor way, Norway. Richardson, J. & Mackay, R. 1991. Lake outlets and the distribution of filter feeders: an assessment of hypotheses. Oikos 62,370-380. Sand, K. 1997. Life cycle studies of Ephemeroptera and Plecoptera in a Norwegian mountain area, 0vre Heimdalen. 26 pp. Thesis, Zoological Museum, University of Oslo, Norway. Solem 1. O. & Gullefors, B. 1995. Trichoptera, caddis flies. Pp. 223-255 in Nillson, A.(ed). The Aquatic Insects of North Europe. Apollo Books, Stenstrup, Denmark. Solem, J. O. 1985. Distribution and biology of caddis flies (Trichoptera) in Dovrefjell mountains. Cen tral Norway. Fauna norv. Ser. B. 32, 62-79. Sweeney, B. W. 1984. Factors influencing life-history patterns in aquatic insects. Pp. 56-100 in Resh, V. H. & RosenbergD. M. (eds.). The Ecology ofAqua tic Insects. Praeger, New York. Szczesny, B. 1986. Caddisflies (Trichoptera) of run ning waters in the Polish North Carapathians. Acta Zool. Cracov. 29, 501-586. Tachet, H. 1967. Quelques aspects du cycle biologique de Plectrocnemia conspersa (Curtis, 1893) (Tricho ptera, Polycentropodidae). Annls. Limn. 3, 177-184. Townsend, C. R. & Hildrew, A. G. 1979. Resource par titioning by two freshwater invertebrate predators with contrasting foraging strategies. J. Anim. Ecol. 48, 909-920. Ulfstrand, S. 1968. Life cycle of benthic insects in Lapland streams (Ephemeroptera, Plecoptera, Tri choptera and Diptera Simuliidae). Oikos Suppl. 10, 1-120. Vik, R. 1978. The lake, 0vre Heimdalsvatn, a subalpine freshwater ecosystem. Holarc. Ecol. I, 81-320. Ward, J. V. & Stanford, 1. A. 1982. Thermal responses in the evolutionary ecology ofaquatic insects. Ann. 124 Norw. J. Entomol. 47, 125-130.2000 Individual variation in behavioural attraction and corresponding olfactory receptor capability to enantiomers of the pheromone ipsdienol in the pine engraver Ips pini Bjorn Age Tommeras Tommenls, B.A. 2000. Individual variation in behavioural attraction and corresponding olfactory receptor capability to enantiomers ofthe pheromone ipsdienol in the pine engraver Jps pini. Norw. J. Entomol. 47, 125-130. Individual variation in pheromone production and behavioural responses elicited by pheromone compounds are shown in bark beetles as well as species of other insect groups. The aim of the present study was to elucidate whether the individual variations are reflected at the level ofolfactory receptor neurons. In the bark beetle Jps pini (Col., Curculionidae, Scolytinae) the relationship be tween individual variation in behavioural attraction and olfactory receptor capability to the enantiomers ofipsdienol are compared. The method applied was recording electroantennograms, EAG. Responses were recorded to 0-25% and 90-100% (-)ipsdienol and no differences in the elicited EAG-responses were discovered between the individuals of the two potentially most extreme behavioural groups. Consequently, the different behavioural responses ofindividuals seem not to be based on variations ofthe olfactory receptor neurons. The behavioural variation seems to be connected to integration on higher level in the central nervous system. Key words: Jps pini, EAG, pheromones, individual variation, olfactory receptor neurons Bjorn Age Tommeras, Norwegian Institute for Nature Research (NINA), Tungasletta 2, N-7485 Trondheim, Norway. E-mail: [email protected]./lO INTRODUCTION Schlyter & Birgersson (1989) reported that the coefficient of variation of the production of phe Intraspecific communication by chemical signals, romone in bark beetles often turned out to be close defined as pheromones (Karlson & Liischer 1959), to 100%. The ratio ofvariation using enantiomers is well studied in species of several insect fami of compounds was lower, approximately 25%. lies. In solitary species, simple pheromone mix Variation in behavioural responses occurred in all tures are used for partner attraction, whereas in tests, but was not easily quantified. It is not well social insects complex mixtures from numerous documented whether the olfactory receptor level glands serve various intraspecific functions. Al or higher levels in the nervous system are respon though not social, numerous individuals ofthe bark sible for the observed individual variation in be beetles of the subfamily Scolytinae live together havioural responses to pheromones. on the same host tree. Several species show mass attack and are considered pest insects. Their chemi In the pine engraver, Ips pini (Say) interpopula cal ecology is rather well studied. Individual vari tional variation in production and behavioural re ation is shown both intra- and interpopulationally. sponse to pheromones is well documented. This This variation relates to pheromone production and is due to the utilisation of different enantiomers release of pheromones, as well as to the behav ofthe pheromone ipsdienol (Birch et al. 1980, La ioural response to pheromones (cf. Byers 1989). nier et aI1980). Electrophysiology, electro-anten nograms (EAG) and single cell recordings, has 125 T0mmeras: Individual variation in olfactory receptor capability in Ips pini been applied in order to detect differences between days. The synthetic enatiomers of ipsdienol were populations in the ability ofthe olfactory receptor kindly provided by Firmenich Company, Switzer neurons to perceive the pheromones. By means of land, and were synthesised according to Ohloff & EAG, Angst & Lanier (1979) was not able to find Giersch (1977). such differences, but the use of single cell record ings indicated differences in the number of iden Preparation and electrophysiological recordings tified receptor neurons detecting the two The preparation ofliving insects of1. pini for elec enantiomers ofipsdienol (Mustaparta et al. 1985). trophysiological recordings followed standard The aim of the present study was to test if indi preparation technique on plexiglas holders de vidually observed behavioural responses of the scribed previously (Angst & Lanier 1979, Tom bark beetle, Ips pini (western population) to the men'ls et al. 1993). Beetles were immobilised in a pheromone (-)ipsdienol (Birch et al. 1980) corre plexiglas holder, and antennal movement was pre sponded to differences in the olfactory receptor vented by placing the antenna onto wax. The re neurons ofthe individual beetles. The method used cording glass capillary electrode filled with insect to study the olfactory abilities of the antenna to ringer solution was inserted into the distal part of perceive the pheromone compounds was record antenna. The indifferent glass electrode was placed ing electro-antennograms (EAG). This is the only in the mouth ofthe insects. possible method when studying relationships be tween behavioural responses and the olfactory Experimental procedure receptor abilities in the same individual. Synthetic compounds from a «syringe olfactom eter» (Kafka 1970) were tested by the stimulation MATERIALS AND METHODS procedure described by Mustaparta et al. (1980). Insects and pheromone substances A syringe containing 0.0 I, 0.1,...,100llg of the compound inserted onto a filter paper was the The insect material, adult beetles of1. pini, origi odour source for stimulation. The synthetic com nated from California, USA. They consisted offive pounds were tested in double series, independently groups which had been collected from traps baited made syringe set A and B, at five concentration with mixtures with different ratios of(+) and (-)ips steps, from the lowest to the highest. Before and dienol, 0%, 25%, 50%, 90%, 100% of the (-)en after a test program a syringe without compounds antiomer. The beetles were transported to Norway was used as pure air stimulation for control as well by air immediately after trapping. The insects were as a syringe with a small piece of bark to check kept in the laboratory at 4-5°C for maximum 20 stable recording conditions. Baseline and EAG Table 1. The successful EAG recordings grouped after sex and previ ous behavioural response to (+) and (-)ipsdienol of the individual bee tles of I. pini. % of (-)ipsdienol behavioural responses in the traps 99 dd Sum 0 1 3 4 25 2 3 5 50 3 3 6 90 0 2 2 100 5 7 12 Log 6 - 6 Random 6 4 10 126 Norw. J. Entomol. 47, 125-130.2000 amplitudes were continuously visualised on an amplitudes (varying from O,75mV to 2,3mV) were oscilloscope and stored on tape. normalised and expressed as a percentage of the mean response of the two syringes containing Evaluation of data 100(g (-)ipsdienol. The amplitude ofEAG responses was determined as the maximum ofnegative deflection ofthe DC RESULTS AND DISCUSSION potential between the electrodes during stimula Successful EAG recordings were obtained from tion. In order to allow comparison ofthe EAG re 45 individuals as shown in Table 1. There were no sponses obtained in different individuals, the EAG differences between females and males as reported Jps pini individuals attracted to 0-25% (-)ipsdienol 120 Syringe set A T • (-)ipsdienol 100 ~ 0 c 80 T/l(+)iPSdienOI GI III . ~~~ C ----.-/1~ 0 T Co 60 III T~I· ~~~-~~~- GI T c::: ~ 40 1---·---::::-1 w 20 0 0.01 0.1 10 100 Dose: Ilg per test syringe Jps pini individuals attracted to 90-100% (-)ipsdienol 120 Syringe set A -. (-)ipsdienol ~ 100 c T~f (+)ipsdienol GI III 80 C 0 T -----:. ,,' Co III 60 ·--",/1 c:::GI ~ 40 ~C- w 20 0 0.01 0.1 10 100 Dose: Ilg per test syringe Figure 1. Dose-response relationships for electroantennogram (EAG) responses elicited of the two enantiomers of ipsdienol from syringe set A. Top: Individuals of Ips pini which previously showed be havioural attraction to 0 or 25% (-) ipsdienol (N=9). Below: Individuals which showed behavioural at traction to 90 or 100% (-) ipsdienol (N=14). Bars refer to standard deviation (SO). 127 T0mmeras: Individual variation in olfactory receptor capability in Ips pini previously on bark beetles (Angst & Lanier 1979, antiomer ofthe pheromone ipsdienol (Figure 1 and Mustaparta et a1. 1979, 1980, T0mmeras 1985, 2). To secure a more reliable dose-response rela T0mmeras & Mustaparta 1989). tionship the two independently made stimulation series were kept separately (syringe set A and B). To allow statistical analysis, the data from the EAG Figure 1 shows the results obtained by syringe sti recordings were grouped. The 9 individuals which mulation of serie A while Figure 2 shows corre in behavioural tests responded to either 0 or 25% sponding data obtained by using serie B as stimu (-)ipsdienol are compared to the 14 individuals lation source. No differences in the elicited EAG which responded to either 90 or 100% ofthe (-)en responses were seen from the recordings when 'ps pini individuals attracted to 0-25% (-)ipsdienol 120 Syringe set B ';fl. 100 =: _ /1 (-)ipsdienol ::: 80 T V -f (+);psdlenol s::: o a- 60 ~ 1~------1--_1------"" ~ 40 1.. "'1 w ------1"'" 20 o I I I I I I 0.01 0.1 1 10 100 Dose: Ilg per test syringe 'ps pini individuals attracted to 90-100% (-)ipsdienol 120 Syringe set B ';fl. 100 T (-)ipsdienol .=: 3l 80 T ,,,1 (+)ipsdienol s::: o T "", a- 60 ~ T_------J[--""--~ ~ 40 w 20 O I , 0.01 0.1 1 10 100 Dose: Ilg per test syringe Figure 2. Dose-response relationships for electroantennogram (EAG) responses elicited of the two enantiomers of ipsdienol from syringe set B. Top: Individuals of 'ps pini which previously showed be havioural attraction to 0 or 25% (-) ipsdienol (N=9). Below: Individuals which showed behavioural at traction to 90 or 100% (-) ipsdienol (N=14). Bars refer to standard deviation (SO). 128 Norw. J. Entomol. 47, 125-130.2000 comparing the individuals from the two potentially References most extreme groups according to the behavioural Angst, M. E. & Lanier, G. N. 1979. Electroantennogram responses to mixtures of(+) and (-)ipsdienol. responses oftwo populations of[ps pini (Coleoptera: Scolytidae) to insect-produced and host-tree com In the present study variation on the olfactory pounds. 1. Chem. Ecol. 5, 131-140. receptor level between individuals with different Birch, M. c., Light, D. M., Wood, D. L., Browne, L. E., behavioural responses was not found. This means Silverstein, R. M., Bergot, B. 1., Ohloff, G., West, that the property ofthe olfactory receptor neurons 1. R. & Young, J. C. 1980. Pheromonal attraction of I. pini seems not to be the basis for the varia and allomonal interruption ofIps pini in California tion in observed behaviour ofindividuals aroused by the enantiomers of ipsdienol. J. Chem. Ecol. 6, by the two enantiomers ofipsdienol. The basis for 703-717. the behavioural variation of individuals seems to Byers, J. A. 1989. Chemical ecology of bark beetles. Experientia 45, 271-283. be different integration on a higher level in the Kafka, W. A. 1970. Molekulare Wechselwirkungen bei central nervous system. der Erregung einzelner Riechzellen. Z. Vergl. It could be argued that the bark beetle species Physiol. 70,105-143. might possess an intermediate type of olfactory Karlson, P. & Uischer, M. 1959. Pheromone. Ein receptor cell for receiving e.g. blends of the two Nomenklatur-vorslag fUr eine Wirkstoffklasse. Naturwissen. 46, 63-64. ipsdienol enantiomers, and this would hide varia Lanier, G.N, Classon, A., Stewart, T., Piston, J.J. & tions that could be discovered by the use of the Silverstein, R.M. 1980. [ps pini: The basis for EAG technique. However, it has previously been interpopulational differences in pheromone biology. shown that the receptor cells are types responding 1. Chem. Ecol. 6,677-687. to either (+) or (-) ipsdienol, and that there are no Mustaparta, H., Angst, M. E. & Lanier, G. N. 1979. intermediate receptor types present in bark bee Specialization of olfactory cells to insect- and host tles (Mustaparta et al. 1985). produced volatiles in the bark beetle Ips pini (Say). 1. Chem. Ecol. 5,109-123. Evaluation of individual differences on the level Mustaparta, H., Angst, M. E. & Lanier, G. N. 1980. of olfactory receptor neurons is in general diffi Receptor discrimination of enantiomers of the ag cult due to differences in recording conditions and gregation pheromone ipsdienol, in two species of the physiological status of the insect from one re Ips. J. Chem. Ecol. 6, 689-701. cording to another. In addition the EAG technique Mustaparta, H., Tommenls, B. A. & Lanier, G. N. 1985. is considered quite coarse as a basis for evalua Pheromone receptor cell specificity in interpopu lational hybrids ofIps pini (Coleoptera: Scolytidae). tion of the importance of chemical agents for the 1. Chem. Ecol. 11,999-1007. insects. These factors are probably not so impor Ohloff, G. & Giersch, W. 1977. Access to optically ac tant uncertainties in the present study when only tive ipsdienol from verbenone. Helve. Chim. Acta testing the relative differences between EAG re 60,1496-1500. sponses elicited of two enantiomers of the main Schlyter, F. & Birgersson, G. 1989. Individual varia pheromone compound ipsdienol. In addition the tion in bark beetle and moth pheromones - a com recording conditions and the physiological status parision and an evolutionary background. Holarc. have been monitored using control stimulations Ecol. 12,457-465. during each test ofthe insects. Tommenls, B. A. 1985. Specialization of the olfactory receptor cells in the bark beetle [ps typographus and its predator Thanasimus fonnicarius to bark beetle Acknowledgements. I especially acknowledge Steven pheromones and host tree volatiles. J. Comp. Phy Seybold and David Wood, University of California, siol. A 157, 335-341. Berkeley, for perfonning the behavioural selection tests Tommeras, B. A. & Mustaparta, H. 1987. Chemorecep of the bark beetle [ps pini and organise the supply of tion ofhost volatiles in the bark beetle [ps typogra living beetles to the laboratory in Norway. I also thank phus. J. Comp. PhysioL A 161, 705-710. Prof. Hanna Mustaparta at Department of Zoology, Tommeriis, B.A. & Mustaparta, H. 1989. Single cell NTNU, head of the laboratory where the experiments responses to pheromones, host and non-host vola were carried out. tiles in the ambrosia beetle Trypodendron lineatum. 129 T0mmeras: Individual variation in olfactory receptor capability in Ips pini Ent. Exp.Appl. 52, 141-148. Tommenls, B. A., Wibe, A., Nilssen, A. & Anderson, J. R. 1993. The olfactory response ofthe reindeer nose bot fly, Cephenemyia trompe (Oestridae), to com ponents from interdigital pheromone gland and urine from the host reindeer, Rangifer tarandus. Chemo ecol. 4, 115-119. Received 29 September 1999, accepted 17 April 2000 130 Norw. J. Entomol. 47, 131-136.2000 A new species of Physoneura Ferrington et Scether, 1995, from Ecuador (Chironomidae, Orthocladiinae) Elisabelh Slur & Trond Andersen Stur, E. & Andersen, T. 2000. A new species ofPhysoneura Ferrington et Srether, 1995, from Ecua dor (Chironomidae, Orthocladiinae). Norw. 1. Entomol. 47,131-136. Physoneura paulseni sp. n. is described and figured as male imago based on specimens collected in the Chimborazo Province in Ecuador. The genus Physoneura Ferrington et Srether, 1995, originally described based on two species from Chile and Argentina, now includes three species from South America. A key to the male imagines is given. Key words: Chironomidae, Orthocladiinae, Physoneura, new species, Ecuador Elisabeth Stur & Trond Andersen, Museum ofZoology, University ofBergen, Museplass 3, N-5007 Bergen, Norway. INTRODUCTION out by Ferrington & Srether (1995), knowledge of the females and the immatures is necessary to place The genus Physoneura was erected by Ferrington the genus with certainty. & Srether (1995) for two Orthoc1adiinae species, P nigroflava (Edwards, 1931) and P. minuscula Ofthe two previously described species, P nigro (Edwards, 1931) from Patagonia and Southern flava (Edwards) is known from Bariloche in Ter Chile. Edwards (1931) placed the two species in ritory Rio Negro in Argentina, while P. minuscula the genus Spaniotoma Philippi, subgenus Smittia (Edwards) was collected in Castro, on the Island Holmgren. The two species show similarities to ofChiloe in southern Chile. Both localities are in several semi-terrestrial genera like Pseudosmittia Patagonia south of 30° south latitude. The newly Goethgebuer and Colosmittia Srether et Andersen. described species was collected in the Chimborazo Brundin (1956) discussed the systematic position Province in Ecuador, where it was taken close to of the two species and doubted they belonged in permanent snow beds at an altitude ofabout 4.800 Pseudosmittia. When erecting the genus Ferring meter. The species thus extends the known range ton & Srether (1995) placed Physoneura in the of the genus north to the central Andes, an area Corynoneura group of genera together with which also has a predominately austral fauna (see Corynoneura Winnertz, Thienemaniella Kieffer e.g. Humphries & Parenti 1999). and Tempesquitoneura Epler et de la Rosa, mainly based on the subapical inflation ofRI and the api METHODS cal inflation of R4+5. The genus also has a small, but distinct keel on the front trochanter and an The material examined was mounted on slides fol oblique tibial comb, which are characters shared lowing the procedure outlined by Srether (1969). with Corynoneura. But Physoneura differs from The general terminology and abbreviations follow the other three genera, particularly in the shape of Srether (1980). The measurements are given as a the phallapodeme, which has a blunt, rounded range followed by the mean. aedeagal lobe, in contrast to the pointed and The holotype and 9 paratypes ofPhysoneura paul strongly recurved aedeagallobe usually found in seni sp. n. are deposited in the Museum of Zoo the Corynoneura group ofgenera. Thus, as pointed logy, Bergen, Norway (ZMBN); 2 paratypes are 131 Stur & Andersen: A new Physoneurafrom Ecuador (Chironomidae, Orthocladiinae) deposited in the Zoologische Staatssammlung, Etymology: Named after Frederik A. Paulsen, for Munich, Germany (ZSM); 2 paratypes in Cole<;ao giving chironomid material from exotic places to Entomologica, lnstituto Oswaldo Cruz, Rio de Ja Bergen Museum. neiro, Brazil (FIOCRUZ); and 2 paratypes in Mu seo Ecuatoriano de Ciencias Naturales, Quito, Description Ecuador (MECN). Male imago (n = 10, ifnot otherwise stated). - To tal length 1.62-1.90, 1.74 mm. Wing length 1.25 SYSTEMATICS 1.35, 1.30 mm. Total length / wing length 1.27 Physoneura Ferrington et Scether, 1995, 1.40,1.32. Wing length/length ofprofemur 3.83 emended 4.16,3.99. Colouration dark brown. Generic description as in Ferrington & Sa:ther Head (Figure 1). Antenna as in Figure 3; AR 0.24 (1995), except stemapodeme without or with dis 0.29,0.27; ultimate flagellomere 124-152, 135 Jlm tinct oral projections. long. Postorbitals 0-2, 2. Clypeus with 4-6, 6 se tae. Cibarial pump, tentorium and stipes as in Fig Key to male imagines of Physoneura ure 2. Tentorium 106-117, III (5) Jlm long; 10 Ferrington et Scether, 1995 13, 12 (5) Jlm wide at sieve pore, 10-14, 13 (5) 1. Small species, wing length about 0.75 mm; Jlm wide at posterior tentorial pit; stipes 88-98, ~+5 ending above apex of Cu 1; abdominal 96 (6) Jlm long; 23-32, 26 (6) Jlm wide. Palp seg segments 1-3 yellowish. Chile. (Ferrington & mentslengthsinJlm: 12-16,15; 18-27,24;37-41, Sa:ther 1995: Figures 6-10)... P minuscula 39; 41-49, 44; 51-74, 59. Third palp segment with (Edwards) 1 sensilla clavata subapically, 10-12, 12 Ilm long. Larger species, wing length> 1.0 mm; ~+5 Thorax (Figure 5). Antepronotum with I seta. ending proximal to apex ofCu1; colouration Dorsocentrals 2-4, 3; prealars 2. Scutellum with 2 uniformly dark brown to black 2 setae. 2. Wing length about 1.04 mm; transverse ster Wing (Figure 4). VR 1.36-1.47, 1.40. False vein napodeme without oral projections; inferior 412-488, 449 Jlm long. Brachiolum with I seta, volsella weak. Argentina. (Ferrington & Sa: other veins bare. Squama bare. ther 1995: Figures 1-5) P nigroflava Legs. Dorsal keel of front trochanter as in Figure (Edwards) 7. Spur offront tibia 29-36,33 Jlm long; spurs of Wing length about 1.30 mm; transverse ster middle tibia 14-18, 16 Ilm and 11-15, 13 Jlm long; napodeme with strong, triangular oral pro of hind tibia 30-37, 33 Jlm and 17-25,21 Ilm. jections; inferior volsella relatively well de Width at apex offront tibia 22-28,24 Ilm; of mid veloped, broadly rounded to subtriangular. dle tibia 21-23, 22 Jlm; ofhind tibia 26-30,28 Jlm. Ecuador. (Figures 1-10).. P paulseni sp. n. Hind tibial comb (Figure 6) with 9-12, 10 setae; shortest setae 15-21, 18 Jlm long; longest setae 25-30,28 Ilm long. Lengths and proportion oflegs Physoneura paulseni new species are given in Table I and 2. (Figures 1 - 10) Hypopygium (Figures 8-10). Anal point not dis Holotype: male, Ecuador: Chimborazo Province: tinctly set off, with 1-4,3 weak setae; laterostemite Refugio Hermanos Carrel, 4.800 m a. s. I., 9. Feb IX with 1-3,2 setae. Phallapodeme 37-43, 411lm ruary 1997, picked from in between hairs on com long. Transverse stemapodeme arcuate, with dis posite flower, 1. Skartveit leg. (ZMBN No.: 279). tinct, triangular oral projections; 51-59, 57 Ilm long. Gonocoxite 102-113, 106 Ilm long; inferior Paratypes: 15 males, as holotype. volsella relatively well developed, broadly roun Diagnostic characters: See key. ded to subtriangular, 33-41, 38 Ilm long, 11-17, 132 Norw. J. Entomol. 47, 131-136. 2000 Table 1. Lengths (in Ilm) of legs for males of Physoneura paulseni sp. n. fe ti tal ta2 ta, ta. ta5 P, 304-344, 320 388-444, 404 172-192,180 96-116, 104 68-80,72 32-40,36 24-28,26 P2 384-412, 399 332-352, 344 140-160, 148 84-116,99 56-76,70 32-36,33 24-32,28 P3 364-408, 391 380-424, 396 192-224, 205 92-116, 105 100-112,105 36-44,38 28-36,29 Table 2. Proportions of legs for males of Physoneura paulseni sp. n. LR BV SV BR P, 0.42-0.46, 0.45 3.71-4.04,3.88 3.94-4.23, 4.04 2.14-2.78,2.35 P2 0.41-0.47,0.43 3.54-4.28, 3.93 4.65-5.26, 5.02 2.27-3.03,2.54 P3 0.45-0.55, 0.52 3.41-3.69, 3.57 3.61-4.39,3.86 2.90-3.75,3.28 Figures 1-3. Physoneura paulseni sp. n., male imago. 1, head; 2, cibarial pump, tentorium and stipes; 3, antenna. 133 Stur & Andersen: A new Physoneurajinm Ecuador (Chironomidae, Orthocladiinae) 4 5 6 Figures 4-6. Physoneura paulseni sp, n., male imago. 4, wing; 5, thorax; 6, tibial comb. 134 Norw. J. Entomol. 47, 131-136.2000 7 ,. ",'" ," " ; J ,l , = j1/"':. \'. ;'.'.'~. I' 'i\:~f '\:,.. :.:.,.h" ." 9 10 Figures 7-10. Physoneura paulseni sp. n., male imago. 7, trochanter of front leg; 8, hypopygium with tergite IX removed, left dorsal aspect, right ventral aspect; 9, gonocoxite and inferior volsella, ventral aspect; 10, tergite IX and dorsal aspect of left gonocoxite and gonostylus. 135 Stur & Andersen: A new Physoneurafrom Ecuador (Chironomidae, Orthocladiinae) 13 Ilm wide, widest at 0.67-0.81,0.72 of length, ending at 0.83-0.89, 0.86 of gonocoxite length. Gonostylus 43-48, 45 Ilm long; megaseta 6-9, 8 Ilm long. HR 2.06-2.65, 2.32; HV 3.71-4.18,3.90. Biology: The species was collected close to a small, slow flowing, shallow stream with substrate of stones, pebbles and gravel. The locality was situated approximately lOO meters below perma nent snow beds at 4.800 m altitude in an area with sparse vegetation. The midges were found sitting in between the hairs on the stem and leaves of the Asteracea eulcilium nivalis Kunth. Acknowledgements. We are indebted to Or. John Skartveit, Museum of Zoology, University of Bergen, for providing us with the Orthocladiinae material he col lected in Ecuador. Thanks are also due to professor Goran Hogstedt, Department ofZoology, University ofBergen, for all help and support, to Paul Ramsay, University of Plymouth and other members of TAXACOM, for iden tifying Cu/citium nivalis Kunth, and to Gladys Ramirez, Museum of Zoology, University of Bergen, for making the slide preparations. Permission to collect in Ecuador was given by INEFAN, Quito. The study ofSouth Ameri can Orthocladiinae is funded through the Norwegian Research Board (NF) project no.: 121975/720. References Brundin, L. 1956. Zur Systematik der Orthocladiinae (Dipt. Chironomidae). lnst. Freshw. Res., Drotning holm, Rapp. 37, 5-185. Edwards, F. W. 1931. Chironomidae. Pp. 233-331, in Diptera of Patagonia and South Chile. Part n. Fas cicle 5. Trustees of the British Museum, London. Ferrington, L. C. & Slether, O. A. 1995. Physoneura, a new genus of Orthocladiinae from Patagonia and South Chile (Diptera: Chironomidae). Aquat. Ins. 17,57-63. Humphries, C. 1. & Parenti, L. R. 1999. Cladistic Bio geography: interpreting patterns of plant and ani mal distributions. Second edition. 187 pp. Oxford Univ. Press, Oxford. Slether, O. A. 1969. Some Nearctic Podonominae, Dia mesinae, and Orthocladiinae (Diptera: Chirono midae). Bull. Fish. Res. Bd Can. 170, 1-154. Slether, O. A. 1980. Glossary of chironomid morphol ogy terminology (Diptera: Chironomidae). Ent. scand., Suppl. 14, 1-51. Received 31 July 2000, accepted 22 September2000 136 Norw. J. Enlomol. 47, 137-147. 2000 Description of females of three species of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) Lene Martinsen & Geir E. E. Soli Martinsen, L. & Soli, G. E. E. 2000. Description of females of three species of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae). NOIw. J. Entomol. 47,137-147. Descriptions ofthe females ofEctrepesthoneura hirta, E. colyeri and E. pubescens are given, based on material from two Norwegian localities. The studied specimens were hatched from deacying wood and collected by use of Malaise traps. The three species can only be separated on differences in the outline ofthe terminalia, especially in the shape ofthe gonapophysis 9. Apreliminary key for females of the studied species is given, together with detailed figures of the terminalia. Key words: Mycetophilidae, fungus gnats, Ectrepesthoneura, female, description. Lene Martinsen & Geir E. E. Soli, Zoological museum, University of Oslo, Sars gate I, N-0562 Oslo. INTRODUCTION British material studied by him, belonged to E. hirta, and included a very brief description, with The genus Ectrepesthoneura Enderlein, 1911 be out figures, of these females and a second female longs to the family Mycetophilidae in the super tentatively ascribed E. pubescens. The female of family Sciaroidea. The genus is commonly as E. hirta has been figured, but not thoroughly de cribed the tribe Leiini (e.g. Edwards 1925, Soli scribed by Soli (1997: Fig. 42A, B). 1997), but in several respects it takes an intermedi ate position between the two tribes Leiini and Gno Chandler (1980) discussed the relationship be ristini. Tuomikoski (1966) and Vaisanen (1986) tween Ectrepesthoneura and Tetragoneura Win have both argued for its inclusion in the latter. nertz, 1846, by American authors usually regarded as synonyms (e.g. Vockeroth 1981), and suggested Ectrepesthoneura is known from the Holarctic that some ofthe ten Nearctic species described as region only, with 12 species from the Palaearctic Tetragoneura should be transferred to Ectrepestho and 3 from the Nearctic subregions (Table 1); two neura. These two genera have commonly been more species have been described from Baltic/ regarded as sistergroups (e.g. Soli 1997), but in a German amber (Eocene/Oligocene), E. magnifica recent paper, Chandler (1999) suggests that Ectre (Meunier, 1904) and E. rottensis Statz, 1944 pesthoneura is paraphyletic and that new genera (Evenhuis 1994). A revision of the European and will be recognized. eastern Nearctic species ofEctrepesthoneura, in cluding an identification key for males, are given Hopefully, more thorough studies offemale Ectre by Chandler (1980). pesthoneura will not only prove valuable in the identification of species, but may also reveal new As for the majority of species ofMycetophilidae, characters that can be valuable in future studies to most females ofEctrepesthoneura have never been solve the phylogenetic relationship ofEctrepestho described, or the descriptions are inadequate, with neura and other enigmatic genera within Myceto out detailed illustrations of the terminalia (e.g. philidae. Caspers 1991; Chandler 1980; Zaitzev 1984). Chandler (1980) assumed that most females in the 137 Martinsen & Soli: Description offemales of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) second segment of palpus, the palpomere ratios Table 1. Extant species of Ectrepesthoneura En derlein, 1911. are given as the length of the first segment to the length of third, fourth and fifth segment, respec PALAEARCTIC: tively. The length of scutum was measured as a E. bucera Plassmann, 1980 straight line from its anterior to its posterior bor E. chandleri Caspers, 1991 der; and the length ofthorax as a straight line from E. colyeri Chandler, 1980 the anterior border of scutum to the posterior bor E. gracilis Edwards, 1928 der of scutellum. The wing length was measured E. hirta (Winnertz, 1846) from the distal median plate to the apex of wing. E. ledenikiensis Bechev, 1988 Two leg ratios are used in the descriptions, given E. montana Zaitzev, 1984 for fore, mid and hind leg, respectively: SY, the E. nigra Zaitzev, 1984 combined length of femur and tibia to the length E. ovata Ostroverkhova. 1977 of the first tarsomere; and BY, combined length E. pubescens (Zetterstedt, 1860) offemur, tibia and first tarsomere to the combined E. referta Plassmann, 1976 length of tarsomere 2-5 (see Soli 1997). In addi E. tori Zaitzev & 0kland, 1994 tion, the lengths of the tibial spurs are given in relation to the diameter of the tibiae, measured NEARCTIC: apically. The arrangement of the tibial setae and E. bicolor (Coquillett, 1901) bristles is indicated by use ofthe following abbre E. laffooni Chandler, 1980 viations: a, anterior; p, posterior; d, dorsal; v, ven E. canadensis Zaitzev, 1993 tral; ad, anterodorsal; av, anteroventral; pd, post erodorsal; pv, posteroventral (McAlpine 1981). Total length is calculated as the combined length MATERIAL AND METHODS of head, thorax and abdomen. When five or more Material specimens were measured the mean value is given in brackets after the range. The material originates from two different locali ties: Frogn, Akershus, SE Norway, andAlta, Finn The morphological terminology follows Soli (1997). mark, N Norway. The material from Frogn was collected by use ofhatching-traps attached to logs Association of females of decaying wood, and the emerging insects were collected in test tubes. These samples held three As all but one of the samples obtained by hatch species of Ectrepesthoneura, E. colyen', E. hirta ing held just one species, the presence of males and one female that could not be identified. The and females in the same sample was considered a sample from Alta held two species, E. hirta and reliable evidence for the association of the sexes. E. pubescens, and were collected by use of Ma The sample with two species contained E. colyeri laise traps. The specimens were stored in 70 % and an unidentified female. Females ofE. colyeri alcohol, and later several of them were dissected were primarily associated on the basis of another and slide mounted according to the method de sample containing just one species. In the mate scribed by Soli (1997). The material is kept in the rial from Frogn, E. colyeri and E. hirta were found collection of Zoological Museum, University of in two different habitats, giving further support to Oslo, Norway. the association. On the basis of the association of females from Frogn and by use of species keys (Chandler 1980; Hutson et al. 1980), males and Measurements and terminology females of E. hirta, and males of E. pubescens, The arrangement ofthe ocelli is described by two were identified in the material from Alta. Conse ocellar ratios defining the position of the lateral quently the remaining female specimens in the ocellus in relation to the median ocelli and the sample from Alta were considered to be E. pube compound eye. Due to the strong reduction ofthe scens. 138 Norw. J. Entomol. 47, 137-147. 2000 B RI R4 Rs ~~~~.~~~!.~I MI CuA2 '-----cuP c Figure 1. - A. Head of Ectrepesthoneura colyeri Chandler. - B. Face and clypeus of E. hirta (Winnertz). - C. Wing of E. colyeri. Abbreviations: A =anal vein; bc =basicosta; C =costa; c1yp =c1ypeus; CuA1 and CuA2 =anterior branches of cubitus; CuP = posterior branch of cubitus; d m p =distal median plate; fc =face; fr =frons; fr fur =frontal furrow; fr tub =frontal tubercle; h =humeral; lac =lacinia; lin =lingua; Ibl = labellum; Ibr = labrum; M1 and M2 = branches of media; pip = palpomere; R1 = anterior branch of radius; R4 and R5 = posterior banches of radius; Rs = radial sector; Sc = subcosta; ta = anterior transversal; tb = basal transversal. 139 Martinsen & Soli: Description offemales of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) STUDIED FEMALES OF ECTREPESTHO crossvein ta. Median and posterior fork complete; NEURA stem of posterior fork very short. All veins with setae except h, Sc, tb, R4, Rs, stem of M, basal General morphology portion ofMI, M2 and crossvein ta. Coloration. Head dark brown. Palpi and mouth Legs. Hind coxa with one row oflong and several parts yellow. Thorax dark brown. Halters yellow smaller setae posteriorly, and additional group of to whitish. Wings clear without markings; costa setae anteriorly, near apex. Tibia and tarsi densely and radius dark, remaining veins yellowish. Legs clothed with short, unevenly arranged, dark trichia. mainly yellowish, trochanters and sometimes also Fore tibia only with setae apically. the basalmost part of coxae brown. Coxa, tro chanter and femur with pale setae. Tibial spurs Terminalia (Figures 2-5). Pair of broad gono yellowish. Abdominal tergites brown or with yel coxites 8 intimately fused with posterior border low markings. of stemite 8, evenly clothed with small setae and several bristles posteriorly. Pair of weakly sclero Head (Figure lA, B). Three ocelli in linear ar tized, subtriangular gonapophyses 8 located dor rangement on top of head. Vertex and lateral por sally of gonocoxites 8. Median, membranous la tions of frons evenly covered with setae of vari bia located between gonocoxites 8 and gonapo able size. Height ofcompound eye about 0.5 times physes 8. Tergite 8 narrow. Tergite 9 longer than height of head. Eyes with shallow invaginations tergite 8, setose. Pair ofgonocoxites 9 located late above antennal socket; 3-4 small interommatidial rally oftergite 9, densely covered with trichia. Go setae. Frontal furrow usually distinct and complete. napophyses 9 plate-like and membraneous, usu Scape with setae on apical half; pedicel with row ally with longitudinal sclerotized keel. Sperma ofapical setae, one bristle. Flagellum with 14 fla thecal eminense usually distinct, ending in mem gellomeres. Surface ofeach flagellomere with tri braneous area probably representing vestiges of chia and rounded depressions, probably represent sternite 9. Spermathecal ducts weakly sclerotized. ing campaniform sensilla. Fused face and clypeus Tergite 10 narrow, with transverse row of long with numerous setae of variable size. Labrum setae posteriorly, median portion of basal border apically broad. Lingua pointed, with several api prolonged cephalad. Stemite 10 narrow, bare. Cer cal fringes. Palpus with 5 segments: first segment ci two-segmented; second, ovate segment much short; second strongly recuced, hardly traceable; smaller than first segment. third segment thickened at middle, with special ised sensilla along inner surface; fourth attached Key to females of E. coiyeri, E. hirta and to third about 1/3 from it's distal end; fifth segment long and slender. Segment 3-5 ofpalpus with some E.pubescens setae. Lacinia oblong. 1. First segment of cercus produced apically be Thorax. Scutum with pale setae arranged in indis yond the attachment of second segment (Fig tinct, broad rows. Most setae small, but posteriorly ure 2C, D). Wing length equal to or less than 3 some long and pronounced bristles. Scutellum with times as long as profemur colyeri two pairs of bristles, and additional small inter Second segment of cercus attached at the ex mittent setae. Antepronotum and proepisternum treme apex offirst segment. Wing length more evenly covered with setae, some long and pronoun than 3 times as long as profemur 2 ced. Remaining pleural sclerites bare. Anapleural suture complete, declining posteriorly. 2. Viewed laterally, dorsal border of gonocoxites 8 distinctly invaginated (concave) towards apex Wings (Figure 1C). Costa produced beyond tip of (Figure 4C). Gonocoxites 8 with a collection of R5. Sc reaching 0.5 to 0.6 distance to base ofRs. strong apical setae. First segment ofcercus about R4 present, forming small, oblong radial cell. 3 times as long as second pubescens Length of crossvein ta subequal to or longer than radial cell. RI short, 0.8 to 1.6 times as long as Viewed laterally, dorsal border ofgonocoxites 140 Norw. J. Enlomol. 47. 137-147.2000 cere II cere I hyp ------\~ gc 8---H-/fYY stl0 ---\-.-----+t'i--+ B ---spd '------t'-/-+-+--hyp st 8 c D Figure 2. Female terminalia of Ectrepesthoneura colyeri Chandler. - A. Ventral view, sternal parts re moved. - B. Dorsal view, tergal parts removed. - C. Lateral view. - D. Dorsal view. Abbreviations: cere =cercus; gc =gonocoxite; gp =gonapophysis; hyp =hypoproct; sp d =spermathecal duct; st =sternite; tg = tergite. 141 Martinsen & Soli: Description offemales of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) 8 even or slightly convex towards apex (Fig M1. Crossvein ta subequal in length to radial cell, ure 3C). Gonocoxites 8 with setae evenly dis with 2-4 setae apically. R1 somewhat longer than persed. First segment of cercus distinctly less ta and fused with costa for short distance. Median than 3 times as long as second hirta fork starting at, or slightly before, level of R4. Veins M1 and M2 without setae on basal 1/5 and The difference in the ratio used in the first couplet to 1/7, respectively; crossvein ta without setae on separate E. colyeri from the remaining two species basal half. is small, and the character should be used with care. Legs. Mid tibial setae: 4-5 small p apically, 6 pv, 4 Ectrepesthoneura co/yeri Chandler pd, 3 ad, and several additional setae on apical border. Hind tibial setae: 7-10 small p, 15-19 pd, Figures lA, C and 2. 13 ad, and a group ofsmall setae posterior apically. Ectrepesthoneura colyeri Chandler, 1980: 32. Leg ratios: femur to tibia 1.06-1.08 (1.07), 1.19 Studied material. AK Frogn: Danemark (EIS 18), 1.23 (1.21), 0.86-0.90 (0.89); SV 2.81-3.03 (3.93), 999 5 crcr 13 May - 8 July 1997, 2 99 W 8 July 3.00-3.30 (3.16), 3.75-3.96 (3.87); BV 2.91-3.18 26 August 1997 (hatching-traps), B. 0kland & L. (3.05), 3.23-3.48 (3.38), 3.94-4.31 (4.12); spurs Martinsen. 2.2-2.7; 2.6-3.0, 3.11-4.2; 2.5-3.1,3.4-4.0. Description (female) Terminalia. Viewed laterally, gonocoxites 8 evenly Total length 3.52-4.07 (3.69) mm (n=8). rounded. Tergite 8 with one row of setae posteri Coloration. Antennae with scape dark brown, pedi orty and few additional setae laterally. Gonocoxites cel and first flagellomere yellow; remaining flagel 9 plate-like, weakly sclerotized. Gonapophyses 9 lomeres brown, not as dark as scape. Trochanters plate-like with uneven border, weakly sclerotized, dark. Femora slightly darkened basely, very weak with keel about 1/4 from basal border. Stemite 10 lyon fore leg. Hind femur darkened at apical fourth subtriangular. Hypoproct with setae laterally, a few to apical third. Abdominal tergites brown with yel long basally. First segment of cerci produced api low markings: I and 2 with yellow areas laterally; cally beyond attachment ofsecond, and about 2.5 3 and 4 with yellow areas posterolaterally making times as long as this. the brown area triangular-shaped; 5 with a slim Biology. Specimens of E. colyeri were hatched yellow area on hind margin; 6 and 7 completely from a rotten trunk of oak (Quercus robur). brown. Abdominal stemites: 1-4 yellow, 5-7 brown ish. Terminalia brown, except for yellow cerci. Ectrepesthoneura hirta (Winnertz) Head. 0.8-0.9 (0.9) times as long as wide. Lateral Figures 18 and 3. ocellus separated from eye border and median Tetragoneura hirta Winnertz, 1846: 19. Fig. 8. ocellus by 3.2-4.3 (3.6) and 3.2-3.8 (3.5) times its Ectrepesthoneura hirta (Winnertz, 1846), Ender own width, respectively. Frontal furrow distinct lein 1911: 155 and complete. Length of flagellum 1.0-1.1 mm, or about 1.2-1.4 (1.3) times as long as scutum. Studied material. AK Frogn: Smihagen (EIS 28), Flagellomeres 1-13 and 14, 1.4-2.0 and 2.5-3.6 4994 crcr 8 July - 26 August 1997,3 crcr 13 May times as long as wide, respectively. Fused face and - 8 July 1997 (hatching-traps), B. 0k1and & L. clypeus with about 40 setae. Pa1pomere ratios Martinsen; FV Alta: Detsika, Buolamalia (EIS 1:3.1-5.0:3.7-5.3:4.7-8.9. 173), 3 9944 crcr 11 June - 3 July 1995 (Malaise trap, sandy slope), L. O. Hansen & H. Rinden. Thorax. Tota11ength 0.90-0.98 (0.93) mm. Lenght of scutum 0.77-0.84 (0.81) mm. Description (female) Wings. Wing length 2.57-2.78 (2.67) mm. Wing Total length 3.57-4.05 mm (n=4). length to length of R1 6.67-8.19 (7.40). Wing Coloration. Antennae with scape dark brown, pedi length to length of profemur 2.81-3.00 (2.91). cel and base of first flagellomere yellow, remain Costa covering about 0.7 distance between R5 to ing flagellomeres brown. Trochanters, base of 142 Norw. J. Entomol. 47, 137-147.2000 /"""'~gc8 fi"i'I'I"\----"·<;:',.\ \ gp 8 labia A B cere I tglO""""- tg 9 "'-I st 8 gp9 c tg 9 ~ \. ;... Figure 3. Female terminalia of Ectrepesthoneura hirta (Winnertz). - A. Ventral view, sternal parts re moved. - B. Dorsal view, tergal parts removed. - C. Lateral view. - D. Dorsal view. Abbreviations: see Figure 2. 143 Martinsen & SOli: Description offemales of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) femora, extreme apex ofmid femur, apical fourth Biology. Specimens ofE. hirta were hatched from ofhind femur and the basalmost part ofhind coxa deacaying trunks ofash (Fraxinus excelsior). The dark brown. Abdomen uniform dark brown, or species has previously been hatched from rotten tergite 1-5 with yellow bands apically. Terminalia wood ofbeach, pine and Coryolus versicolor, and yellow, except for brown cerci. from fungus encrusted bark (Chandler 1980). Head. 0.7-1.0 times as long as wide. Lateral ocel lus separated from eye border and median ocellus Ectrepesthoneura pubescens (Zetterstedt) by 2.2-2.5 and 2.6-3.1 its own width, respectively. Figure 4. Frontal furrow distinct, with rupture near frontal Sciophila pubescens Zetterstedt, 1860: 6559 tubercle. Several long, pronounced setae posteri Ectrepesthoneura messaurensis Plassmann, 1975: 10. orly on vertex, reaching down to level ofantennal attachment. Length of flagellum 1.04-1.08 mm, Studied material. FV Alta: Detsika, Buolamalia or about 1.1-1.4 times as long as scutum. Flagel (EIS 173), 149914 aa II June - 3 July 1995 lomeres 1-13 and 14,1.6-1.9 and 2.1-3.3 times as (Malaise trap, sandy slope), L. O. Hansen & H. long as wide, respectively. Fused face and clypeus Rinden. with about 50 setae. Palpomere ratios 1:2.5 Description (female) 2.7:2.7-4.0:4.2-6.0. Total length 3.43-3.67 (3.61) mm (n=6). Thora.,. Total length 0.90-1.08 mm. Length ofscu Coloration. Scape dark brown, pedicel and basal tum 0.78-0.93 mm. half or entire first flagellomere yellow. Remain Wings. Wing length 2.62-3.36 mm. Wing length ing flagellomeres brown. Fore and mid coxae to length of RI 6.08-6.60. Wing length to length slightly darkened at extreme base. Trochanters and of profemur 3.19-3.38. Costa covering about 0.7 extreme base of femora dark. Mid femur dark distance between R5 and MI. Crossvein ta longer ened at extreme apex. Hind femur darkened at api than radial cell, with 8 setae. RI short, but longer cal third. Tibial spurs yellowish. Abdominal ter than tao Median fork starting at level ofR4. Basal gites brown. Terminalia light brown. 1/6 ofMI, extreme bases ofM2 and crossvein ta Head. 0.7-0.9 (0.8) times as long as wide. Median without setae. ocellus slightly smaller than laterals. Lateral ocel Legs. Mid tibial setae: 4 p, 6 small pv, 3 pd, 4 ad, lus separated from eye border and median ocellus and several additional setae on apical border. Hind by 2.5-3.3 (2.8) and 2.5-3.7 (3.0) times its own tibial setae: 6-8 small p, 12 pd, I d basally, 14 ad, width, respectively. Frontal furrow nearly distinct, and a group of small setae posterior apically. Leg with rupture near frontal tubercle. Length of ratios: femur to tibia 0.92-1.11, 0.93-1.08, 0.79 flagellum 0.80 mm (n= I), or 1.3 times as long as 0.83; SV 2.60-2.73, 2.87-3.07, 3.44-3.82; BV scutum. Fused face and clypeus with 30-35 setae. 2.35-2.83,2.98-3.35,4.12-4.46; spurs 2.1-2.9; 3.1 Palpomere ratios I : 2.5-2.7 : 2.8-3.6 : 4.4-6.3. 3.3,3.3-4.3; 2.9-3.1, 3.0-3.9. Thorax. Total length 0.73-0.80 (0.74) mm. Length Terminalia. Viewed laterally, gonocoxites 8 evenly of scutum 0.58-0.68 (0.63) mm. rounded. Tergite 8 with some lateral setae. Wings. Wing length 2.16-2.45 (2.31) mm. Wing Gonocoxite 9 plate-like. Gonapophyses 9 with length to length of RI 7.06-9.07 (8.09). Wing well scierotized, curved lateral border; otherwise length to length of profemur 3.35-3.50 (3.43). weakly sclerotized. Vestiges of stemite 9 as very weakly sclerotized rectangular plate. Stemite 10 Costa covering about half or somewhat more of distance from R5 to MI. Crossvein ta subequal in oblong. Hypoproct with several lateral setae, length to radial cell, with 5-6 setae. RI somewhat basalmost very long. First segment of cercus cy longer than tao Median fork starting at, or slightly lindrical, 2.1-2.4 times as long as second; second before level of R4. Basal fourth of ta, extreme segment ovate, situated at extreme apex of first. base ofMI and basal third ofM2 without setae. 144 Norw. J. Entomol. 47, 137-147.2000 ~ I cerc II \. :: cerc I : :','.: st 10 '( " gc 8 I>! hyp ~,\ gp8 tg 8 A B tglO / hyp c Figure 4. Female terminalia of Ectrepesthoneura pubescens (Zetterstedt). - A. Ventral view, sternal parts removed. - B. Dorsal view, tergal parts removed. - C. Lateral view. - D. Dorsal view. Abbreviations: see Figure 2. 145 Martinsen & Soli: Description offemales of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) Legs. Mid tibial setae: 2 small p, 4 small pv, 2 pd, species. Despite being difficult to observe, the 4 ad , and several additional setae on apical bor shape and position of labia seems to be a good der. Hind tibial setae: 10 minute p, 12 pd, 8 ad, character to separate between the species. In E. and a group of small setae posterior apically. Leg colyeri, labia is clearly attached laterally to gona ratios: femur to tibia 1.03-1.08 (1.05), 1.12-1.22 pophysis 8, but not so in any ofthe other observed (1.15), 0.83-0.86 (0.84); SV 2.93-3.33 (3.14), species. E. colyeri also has the broadest labia, 3.24-3.42 (3.33), 3.58-4.05 (3.77); BV 3.07-3.35 while in E. pubescens labia can only be observed (3.26),3.48-4.16 (3.72), 4.10-4.50 (4.30); spurs as a transverse, narrow bridge. The gonocoxites 9 2.2-2.3; 2.8-3.0, 3.6-3.7; 2.4-2.6, 3.1-3.4. vary slightly in shape and attachment to tergite 9 in the studied species, but E. pubescens is the only Terminalia. Viewed laterally, gonocoxites 8 ta species with trichia basally on gonapophysis 9. pered towards apex, with numerous apical setae giving brushy appearance. Tergite 8 with 2 small, Females of E. pubescens are in general smaller median setae; additional setae laterally. Gonocox than females ofboth E. colyeri and E. hirta. Body ites 9 rounded, small. Gonapophysis 9 curved, size, however, is known to vary considerably plate-like, with trichia. Sternite 10 rectangular within species of Mycetophilidae (e.g. Vaisanen with pointed apex. Hypoproct with setae laterally, 1984, Soli 1997), and can usually not be used as a basalmost very long. First segment ofcercus long, diagnostic character. Colouration and colour pat cylindrical, 2.8-3.3 times as long as second; sec tern represent another type ofcharacter that should ond segment ovate, situated at extreme apex of be used with great care. In some genera, however, first. such characters seem rather constant and appar ently separate well between species. This may also CONCLUSIVE REMARKS be true for E. colyeri and E. hirta, of which the former has brown terminalia with yellowish cerci, In the keys given by Chandler (1980) and Hutson while E. hirta has yellowish terminalia and brown et al. (1980) several non-genital characters are cercI. used in the identification of E. colyeri, E. hirta and E. pubescens. Due to intraspecific variation The single female that could not be identified was these characters were found to be unreliable in hatched from decaying oak wood, together with the present study; in females such variation was E. colyeri. The species differed most pronounced found to be even larger than in males. Hence, only in the large, rounded gonocoxites 9, and in the characters dealing with the terminalia have so far, position ofthe two gonapophyses 9, bordering the proved to separate satisfactorily between females spermathecal ducts medially. Most probably the of the studied species. female belongs to one of the species already re corded from SE Norway, viz. E. tori, E. bucera, In addition to characters used in the key, the char E. referta or E. nigra (see e.g. 0kland & Zaitzev acters that most clearly distinguish between fe 1997). males applies to the chaetotaxy and/or outline of gonocoxites 8, labia, gonocoxites 9, gonapophysis References 9 and the area surrounding the spermathecal emi nence. Except for the outline of gonocoxites 8, Caspers, N. 1991. New and remarkable species of My these characters can only be studied in cleared, cetophiloidea (Diptera, Nematocera) from the Me diterranean region. Spixiana 14 (3), 321-338. preferably slide mounted specimens. Chandler, P. J. 1980. The European and eastem Nearctic In E. pubescens the most prominent setae on fungus-gnats in the genus Ectrepesthoneura (Myce gonocoxites 8 are located distally, seemingly tophilidae). Syst. Entama!. 5, 27-41. forming a thin brush; in E. hirta, and most clearly Chandler, P. J. 1999. Creagdhubhia mallochorum gen. and sp. n. (Diptera: Mycetophilidae), a remarkable in E. colyeri, these setae are more evenly disper new Scottish gnat with a discussion of its relation sed. Moreover, in E. pubescens the two gonocoxi ships. Brit. 1. Ent. Nat. Hist, 12, 121-134. tes 8 are more slender than in any of the other Edwards, F. W. 1925. British Fungus-Gnats (Diptera, 146 Norw. J. Entomol. 47, 137-147. 2000 Mycetophilidae), with a revised Generic Classifica tion ofthe Family. Trans. R. ent. Soc. Lond. 1924, 505-670. Evenhuis, N. L. 1994. Catalogue of the fossil flies of the world (Insecta: Diptera). Backhuys Publishers, Leiden. Hutson, A. M., Ackland, D. M., & Kidd, L. N. 1980. Mycetophilidae (Bolitophilinae, Ditomyiinae, Dia docidiinae, Keroplatinae, Sciophilinae and Mano tinae). In M. G. Fitton (Ed.), Handbooks for the identification of British Insects. Vol. IX, Part 3, 1 112 pp. Royal Entomological Society of London, London. McAlpine, 1. F. 1981. Morphology and tenninology adults. Pp. 9-63 in J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. 1. Teskey, J. R. Vockeroth, & D. M. Wood (Eds) Catalog of Nearctic Diptera (Vo!. I). Research Branch Agriculture Canada, Ottawa. 0kland, B. & Zaitzev, A. I. 1997. Mycetophilids (Di ptera, Sciaroidea) from southeastern Norway. Fauna Norv. Ser. B 44 (I): 27-37. Soli, G. E. E. 1997. On the morphology and phylogeny ofMycetophilidae, with a revision ofCoelosia Win nertz (Diptera, Sciaroidea). Ent. scand. Supp!. 50, 1-139. Tuomikoski, R. 1966. On the subfamily Manotinae Edw. (Diptera, Mycetophilidae.). Ann. Ent. Fenn. 32 (3), 211-223. Viiisiinen, R. 1984. A monograph ofthe genus Mycomya Rondani in the Holarctic region (Diptera, Myceto philidae). Acta. Zool. Fenn. 177, 1-346. Viiisiinen, R. 1986. The delimitation ofthe Gnoristinae: criteria for the classification of recent European genera (Diptera, Mycetophilidae). Ann. Zoo!' Fenn, 23, 197-206. Vockeroth, 1. R. 1981. Mycetophilidae. Pp. 223-247 in J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. 1. Teskey, J. R. Vockeroth, & D. M. Wood (Eds) Cata log ofNearctic Diptera (Vo!. I). Research Branch Agriculture Canada, Ottawa. Zaitsev, A. I. 1984. [New and little known species of subfamily Sciophilinae (Diptera, Mycetophilidae)] Vest. Zoo!. 4, 15-18. (In Russian.). Received 26 October 2000, accepted 7 November 2000 147 Short communication Euides speciosa (Boheman, 1845) The record (Homoptera, Delphacidae) in Norway Four males of the planthopper E. speciosa were captured in a malaise-trap situated at AK Oslo: 0stensjovannet (Northern shore) (EIS 28) 1 - 31 Lars Ove Hansen July 1996, leg. Lars Ove Hansen & Morten Falck. The locality is a typical eutrophic freshwater shore dominated by Phragmites communis together with Hansen, L. O. 2000. Euides speciosa (Boheman, 1845) other !imnic shore plants (e.g. Iris, Typhae, Jun (Homoptera, Delphacidae) in Norway. Norw. 1. Ento cus), and some alder forest (Alnus incana) as well. mol. 47, 148. The plant hopper Euides speciosa (Homoptera, Delpha Biology and distribution cidae) is recorded for the first time in Norway. Four E. speciosa is widespread in both !irnnic and marine males were captured at the northern shore ofLake 0sten shores, most abundant on P communis (Jensen-Haa sjovannet, Oslo, July 1996. Notes on distribution and biology are briefly given. rup 1920). The biology seems quite similar to that ofthe two northern European species of Delphax. Key words: Euides speciosa, Homoptera, Delphacidae, E. speciosa is found widespread in Europe, east Lake 0stensjovannet, new to Norway. wards to Kazakhstan (Ossiannilsson 1978). In Sweden it is not rare in the SE parts of the coun Lars Ove Hansen, Zoological Museum, University of Oslo, Sarsgate 1, NO-0562 Oslo. Norway. try, where it is recorded north to Uppland (Up) and Niirke (Nii). It seems scarce but widespread in Denmark, and very rare in Finland. Introduction Acknowledgements. I am indebted to Morten Falck for The Lake 0stensjrJVannnet is situated only 5 km help during the field work, and to 0stensjovannets from the city centre of Oslo. It is a well known Venner (i.e. Friends of0stensjovannet), represented by resource and picnic area, used for many years as a Finn Gulbrandsen, who financed this insect study at Lake 0stensjovannet. Also thanks to Anne Mette Aust study site for the University of Oslo. In 1995 an myr for kindly correcting the English. investigation was initiated to get a more total sur vey of the insect fauna in this area, in particular References the terrestric groups. Until present a total of 1800 insect species are recorded from this Lake area Hansen, L.O. & Falck, M. 2000. Insektfaunaen ved (Hansen & Falck 2000), including the first record 0stensjovannet, Oslo kommune. 75 pp. 0stensjo in Norway of the planthopper Euides speciosa. vannets venner, Oslo. Jensen-Haarup, A.C. 1920. Cikader. Danmarks Fauna The subfamily De1phacinae includes only three ge 24. 189 pp. G.E.C. Gads Forlag, Kobenhavn. nera in N Europe, i.e. Delphax, Conomelus and Ossiannilsson, F. 1978. The Auchenorrhynca (Homo Euides (Ossiannilsson 1978). Euides speciosa, the ptera) of Fennoscandia and Denmark. Fauna Ent. only species occuring in N Europe of this genus, Scand. 7 (1),1-222. resembles Delphax as imago, but they may easily be distinguished by their antennae. Delphax has Received 18 October 2000, the first antennal segment considerably longer than accepted 10 November 2000 the second, while the first segment in Euides is shorter than the second (Jensen-Haarup 1920). 148 Norw. J. Entomol. 47, 149-152.2000 Spiders (Araneae) new to Norway Kjetil Aakra, Arne Bretten & Otto Frengen Aakra, K., Bretten, A. & Frengen, O. 2000. Spiders (Araneae) new to Norway. Norw. J. Entomol. 47,149-152. Haplodrassus umbratilis (L. Koch, 1866), Drassyllus pumilus (c. L. Koch, 1839) Zelotes puritanus Chamberlin, 1922 (Gnaphosidae), Lepthyphantes abiskoensis (Holm, 1945) (Linyphiidae) and Achaearanea riparia (Blackwall, 1834) (Theridiidae) are reported for the first time from Norway. Data on distribution and ecology are provided for each species. Key words: Aranea, new to Norway. Kjetil Aakra & Otto Frengen, Museum o/Natural History and Archaeology, Department o/Natural History, NTNU, N-7491 Trondheim, Norway. Arne Bretten, Aunli, N-7070 Bosberg, Norway. INTRODUCTION Bloksberg (EIS 20), 3-29 June 1995,30'0', 19 (leg. O. Hanssen & J. I. I. Batvik). Moss: Jel0y, Hvit The species reported here were all discovered in tingbukta (EIS 19) 3-30 June 1995, lO' (leg. O. SE Norway. Four ofthe species were first collected Hanssen & G. Hardeng). by the second author (AB) and the fifth by the third author (OF). Additional specimens oftwo of This is a widespread and common species occur these species were discovered by the first author ring north to Uppland in Sweden (Jonsson pers in material originating from a survey of rare in comm.), in southern parts of Finland (Palmgren sect habitats in the Oslofjord region (see Hanssen 1943), southern Great Britain (Roberts 1995), most & Hansen 1998 for description ofthese localities). parts of continental Europe (Grimm 1985) and as The locality at Furuberget, Hedmark is a pine for far east as South Siberia (Mikhailov 1997). For est on calcareous ground with a dense cover of this reason the present discovery is not suprising needles. Bj0rgedalen is a deciduous forest with and H umbratilis is presumably also widespread little or no ground vegetation. in SE Norway. It is mainly found in dry, stony habitats, including heaths (Roberts 1995) and co Abbreviations offaunal provinces follow 0kland niferous forests (Grimm 1985). (1981). Nomenclature is according to Platnick (1998). The material is deposited in the collecti ons ofthe Museum ofNatural History andArchae Drassyllus pumilus (C. L. Koch, 1839) ology, Department ofNatural History, NTNU. All Material: AK Asker: Bj0rkas (EIS 28), 4 June-2 specimens were captured by pitfall traps except July 1995, 10' (leg. L. O. Hansen & O. Hanssen). where noted. HES Hamar, Furuberget (EIS 46), 1-29 May 1993, 40'0' (leg. AB). RESULTS AND DISCUSSION D. pumilus is known from Osterg6tland and Got GNAPHOSIDAE land in Sweden (Jonsson pers comm.), southwest ern parts of Finland (Palmgren 1943, Lethinen et Haplodrassus umbratilis (L. Koch, 1866) al. 1979), but has not been recorded from Great Material: HES Hamar: Furuberget (EIS 46), 1- 29 Britain or Ireland (Locket et al. 1974, van Hels May 1993, 2(Je) (leg. AB). 0 Fredrikstad: Hank0, 149 Aakra, Bretten & Frengen : Spiders (Araneae) new to Norway dingen 1996). It is also found in central and east ther been reported from Finland (Palmgren 1977), em parts ofcontinental Europe (Grimm 1985). The nor Great Britain or Ireland (Locket et al. 1974, species is probably more widespread than current van Helsdingen 1996). There are otherwise only a records suggest, but apparently has a scattered dis few scattered records from continental Europe: tribution in Fennoscandia. D. pumilus is a ther Switzerland and Austria (Thaler 1981, 1997, Mus mophilous spider occurring in various sun-exposed ter 1997), southern Germany (two records each in habitats (Thaler 1981, Grimm 1985). Saxony and Bavaria and one in Thuringia - Grimm 1985, Uhlenhaut 1990, Stumpf & Blick pers. Zelotes puritanus Chamberlin, 1922 comm.), former Czechoslovakia (Miller & Buchar 1977) and Poland (Starega 1972). The record re Material: HES Hamar: Furuberget (EIS 46), 1-29 ported here is the northermost in Europe. Outside May 1993, 2eX], 399 (leg. AB). Europe, Z. puritanus is known from Russia, north Z. puritanus is only known from Sodermanland eastern Siberia (Mikhailov 1997), Mongolia in Sweden (Sandstrom et al. 1994) and has nei- (Marusik & Logunov 1998), Tuva (Marusik et al. ;!•..•• , • ~ ••~._•• , •• ~ 0, B A Figure 1. Zelotes puritanus. Male palpal organ, ventral view (A) and epigyne (B). 150 Norw. J. Entomol. 47, 149-152.2000 2000) and western parts ofNorth-America (Grimm THERIDIIDAE 1985). The distribution of the species is disjunc Achaearanea riparia (Blackwall, 1834) tive (Marusik et al. 2000). Material: (:) Hvaler: Kirk0y, Stakhalden, 4 July Like the preceeding species, Z. puritanus is ther 2000, lcJ, sweep netting (leg. OF). mophilous. In summarising the known occurences The discovery of this species in Norway was ex in Europe, Grimm (1985) mentioned records from pected as it occurs north to Helsingland in Swe heathlands and forest habitats. A more recent find den (L. J. Jonsson pers. comm.) and north to 62°N ing (H. Stumpfpers. comm.) from a steep south in Finland (Palmgren 1974). It is thermo- and pho wards facing limestone slope (45°) in Wiirzburg, tophilous, having been found in overhanging Northern Bavaria, highlights the thermophilous banks, on stony ground, below flat stones and on requirement of this species. European records of stonewalls (Roberts 1995, Palmgren 1974). It is Z. puritanus are from the most continental parts common and widespread in Europe (Heimer & of the region, suggesting that the distribution in Nentwig 1991). Norway is rather limited. The species is generally found at high elevations in continental Europe; at 700-1200 m in Switzerland and Austria (Thaler Acknowledgements. Many thanks to Helmut Stumpf and Theo Blick for information on the habitat and dis 1981), and Asia; at 1700-1800 m in Tuva, Russia tribution ofZ. puritan US, to Lars J. Jonsson for distribu (Marusik & Logunov 1994) and at 1700 m in Mon tional data from Sweden and to Lars Ove Hansen and golia (Marusik & Logunov 1998). At northern lati Oddvar Hanssen who kindly allowed the first author tudes in Europe the species is found at lower el access to the Oslofjord material. evations. Furuberget is very close to the railway and main roads passing through Hamar city and References an anthropogenic mode of dispersal into Norway Grimm, U. 1985. Die Gnaphosidae Mitteleuropas cannot be ruled out. The male palpal organ (Fig (Arachnidae, Araneae). Abh. Naturwiss. Ver. Ham ure lA) and the epigyne (Figure IB) are very dis burg NF 26, 1-318. tinctive. Hanssen, O. & Hansen, L. O. 1998. Verneverdige insekt habitater. Oslofjordomn'ldet. NINA Oppdragsmel L1NYPHIIDAE ding 546, 1-132. Heimer, S. & Nentwig, W. 1991. Spinnen Mitteleuropas. Lepthyphantes abiskoensis (Holm, 1945) Ein Bestimmungsbuch. 543 pp.Verlag Paul Parey. Material: HES Ringsaker: Bj0rgedalen (EIS 46), Helsdingen, P. J. van 1996. The county distribution of 1-29 May 1993, 1<;?(leg.AB). Irish spiders. 92 pp. The Irish Naturalists' Journal. Special Zoological Supplement 1996. Predominantly a northern species reported from Lehtinen, P. T., Koponen, S. & Saaristo, M. 1979. Stud Lule and Tome Lappmark in Sweden (Jonsson ies on the spider fauna ofthe southwestern archipe pers. comm.) and the southern half of Finland lago ofFinland II. Memo. Soc. Fauna Flora Fennica (Palmgren 1976) which has not been found in 55,33-52. Great Britain or Ireland (Locket et al. 1974, van Locket, G. H., Millidge, A. F. & Merrett, P. 1974. Brit Helsdingen 1996) or continental Europe (Heimer ish Spiders Vol. III. 314 pp. Ray Society, London. Marusik, Y. M. & Logunov, D. V. 1994. Gnaphosid spi & Nentwig 1991). According to Mikhailov (1997) ders from Tuva and adjacent territories, Russia (Ara it has been recorded from the Russian plain and nei: Gnaphosidae). Beitr. Araneol. 4, 177-210. as far east as Shakalin. The species is presumably Marusik, Y. M. & Logunov, D. V. 1998. On the spiders widespread in Fennoscandia. The current record (Aranei) collected in central Mongolia during ajoint is the westernmost according to Marusik et al. American-Mongolian-Russian expedition in 1997. (2000). 1. abiskoensis has been found in various Arthropoda Selecta 7, 233-254. types ofvegetation and appears to prefer the lower Marusik, Y. M. , Logunov, D. V. & Koponen, S. 2000. field layers (Palmgren 1976). Spiders of Tuva, South Siberia. 252 pp. Magadan. IBPN FEB RAS. 2000. Mikhailov, K. G. 1997. Catalogue ofthe spiders of the 151 Aakra, Bretten & Frengen : Spiders (Araneae) new to Norway territories of the fonner Soviet Union (Arachnida, Landkreis Hof, Bayern, SE-Deutschland (Arach Aranei). 416 pp. Zoological Museum of the Mos nida: Araneae). Senckenberg. bioI. 70, 179-183. cow State University. Moscow. 1997. Miller, F. & Buchar, J. 1977. Neue spinnenarten aus der Received 28 August 2000, Gattung Zelotes Gistel undHaplodrassus Chamber accepted 28 September 2000 lin (Araneae, Gnaphosidae). Acta Univ. Carol. BioI. 1974,151-171. Muster, C. 1997. Zur Spinnenfauna der Siichsischen Schweiz: Artenspektrum, Phiinologie und Okologie der Lycosidae, Zoridae und Gnaphosidae (Arach nida: Araneae). Faun. Abhandl. 21, 13-52. 0kland, K. 198 I.Inndeling av Norge til bruk ved bio geografiske oppgaver - et revidert Strand system. Fauna (Oslo) 34, 167-178. Palmgren, P. 1943. Die Spinnenfauna Finnlands. II. Pisauridae, Oxyopidae, Salticidae, Clubionidae, Anyphaenidae, Sparassidae, Ctenidae, Drassidae. Acta. Zool. Fenn. 36,1-112. Palmgren, 1974. Die Spinnenfauna Finnlands und Ostfennoskandiens. V. Theridiidae und Nesticidae. Fauna Fennica 26, I-54. Palmgren, P. 1976. Die Spinnenfauna Finnlands und Ostfennoskandiens. VII. Linyphiidae 2. Fauna Fen nica 29, 1- 126. Palmgren, P. 1977. Die Spinnenfauna Finnlands und OstFennoskandiens VIII. Argyronetidae, AgeIeni dae, Hahniidae, Dictynidae, Amaurobiidae, Titano ecidae, Segestridae, PholcidaeAnd Sicariidae. Fau na Fennica 30: I-50. Platnick, N. I. 1998. Advances in spider taxonomy 1992 1995. With redescriptions 1940-1980.976 pp. Ent. Soc. & Amer. Mus Nat. Hist. New York. Roberts, M. J. 1995. Spiders of Britain & Northern Europe. Collins Field Guide. 383 pp. HarperCollins Publishers. England. Sandstrom, J., Viklund, B. & Vikars, L. -0. 1994. Ved insekter och spindlar i Nykopings kommun. Resultat fran en entomologisk inventering av 22 naturom raden. Nykopings kommun, Kommunledningskon toret, Planering. Rapport 1994-9N, 22 pp. Starega, W. 1972. Nowe dla fauny polskii i rzdadsze gatunki pajakow (Aranei) z opisem Lepthyphantes millen' sp. n. Fragm. Faun. Polska Akad. NAUK Inst. Zool. 18, 55-98. Thaler, K. 1981. Bemerkenswerte Spinnenfunde in Nordtirol (Ostereich). Veroff. Mus. Ferdinandeum Innsbruck 6, 105-150. Thaler, K. 1997. Beitriige zur Spinnenfauna von Nord tirol- 3: «Lycosaefonnia» (Agelenidae, Hahniidae, Argyronetidae, Pisauridae, Oxyopidae, Lycosidae) und Gnaphosidae (Arachnida: Araneae). VerOff. Mus. Ferdinandeum Innsbruck 75/76,97-146. Uhlenhaut, H, 1990. Bemerkenswerte Spinnenfunde im 152 Norw. J. Entomol. 47, 153-162.2000 Noteworthy records of spiders (Araneae) from central regions of Norway Kjetil Aakra Aakra, K. 2000. Noteworthy records of spiders (Araneae) from central regions of Norway. Norw. J.Entomol. 47,153-162. New records are presented for a number ofspecies recently discovered in Central Norway. Caviphantes saxetorum (Hull, 1916) (Linyphiidae) and Arctosa stigmosa (Thorell, 1875) (Lycosidae) are new to Fennoscandia while Singa nitidula C. L. Koch, 1844 (Araneidae) and Myrmarachneformicaria (De Geer, 1778) (Salticidae) are new to Norway. Three rare araneids, Araneus nordmanni (Thorell, 1870), Araneus saevus L. Koch, 1872 and Gibbaranea omoeda (Thorell, 1870), have not been recorded from Norway for almost 100 years. The northernmost records in Europe of Anyphaena accentuata (Walckenaer, 1802) (Anyphaenidae), Micaria subopaca Westring, 1861 (Gnaphosidae), Troxochrus nasutus Schenkel, 1925 (Linyphiidae), Tegenaria atrica C. L. Koch, 1843 (Agelenidae) and Dipoena torva (Thorell, 1875) are also given as well as new distributional data on Arctosa cinerea (Fabricius, 1777) (Lycosidae). Key words: Araneae, distribution in Norway. Kjetil Aakra, Museum ofNatural History and Archaeology, Department ofNaturaI History, NTNU, N-7491 Trondheim, Norway. INTRODUCTION Hanssen 1994) and (4) collections made by the author (KA) and Otto Frengen (OF) in spring and Relatively few papers have included data on the summer 2000. The spider material collected by spider fauna of central Norway, i.e. the counties NINA has only been partly incorporated into the ofSor- and Nord-Trondelag (Collett 1876, 1877, reports mentioned above. Storm 1898, Strand 1904a,b, Tambs-Lyche 1940, 1941, Hauge 1972, 1989, Solem & Hauge 1973, Abbreviations of faunal provinces are according Hauge & Furunes 1976, Dolmen 1977). In previ to 0kland (1981), the nomenclature follows Plat 0usy undetermined collections ofspiders from the nick (1998, 2000). The material is deposited in region, the author discovered several species the collections ofthe Museum ofNatural History which are either new to Fennoscandia and/or Nor and Archaeology, Department ofNatural History, way, have not been found for almost 100 years or NTNU, except where otherwise noted. Other ab which have a new northern limit in Europe. breviations used are: PT (pitfall traps), MT (ma laise traps), WT (window traps), ZMO (Zoologi The new records presented here originate from cal Museum, Oslo). four sources: (I) the depository ofthe Museum of Natural History and Archaeology (MNHA), (2) the forest fragementation experiment carried out RESULTS AND DISCUSSION by NINA in the coastal spruce forest at Mosvik AGELENIDAE north ofTrondheim (Tommeriis & Breistein 1995 Tegenaria atrica C. L. Koch, 1843 Tommeriis et al. 2000), (3) the investigation of the riparian evertebrate fauna of Gaula south of Material: STI Trondheim: Lade, 7 April 2000, Trondheim supervised by NINA (Andersen & 19, inside building (leg. U. Mostervik). 153 Aakra: Records of spiders (Araneae) from central Norway This is the northernmost record from Europe. Pre Araneus saevus L. Koch, 1872 viously the species has only been reported from Material: NTI Mosvik: Meltingen (EIS 97), 7 Sep various cities in South Norway: Oslo, Larvik, tember 1996, 1<;.>, MT, coastal spruce forest (Tem Arendal, Kristiansand, Stavanger, Bergen (Hauge meras et al. 2000). 1989). It is known north to Uppland in Sweden (Jonsson pers. comm.) and from Helsinki in Fin This species was previously known from a few land (Palmgren 1977). old records; «Drammen-Ullensakem and «Arnot - Elverum» (Collett 1877). A. saevus is generally a rare species taken as far north as Asele Lappmark ANYPHAENIDAE in Sweden (Jonsson pers. comm.) and found al Anyphaena accentuata (Walckenaer, 1802) most north to the Polar Circle in Finland (Palmgren Material: STI Melhus: Gravrak (EIS 92), 18 April 1974, 1977). It is not known from Great Britain 2000, 3 subadults, handpicked from under bark or Ireland (Locket et al. 1974, van Helsdingen on Alnus incanus and Salix triandra by the river 1996), but is present in central parts of continen Gaula (KA). tal Europe, northern Asia and North-Amerika (Esyunin & Efimik 1996, Mikhailov 1997). The northernmost record ofthis species from Eu rope. A. accentuata is previously known north to This species is another of our canopy spiders Hordaland (Hauge 1989). In Sweden it occurs (Heimer & Nentwig 1991), although the specimens north to Dalarne (Jonsson pers. comm.) while in reported by Tullgren (1952) and Palmgren (1974) Finland it has been found in southern parts of the where found near or on buildings. country (Palmgren 1943). Gibbaranea omoeda (Thorell, 1870) ARANEIDAE Material: NTI Mosvik: Meltingen (EIS 97), 25 Araneus nordmanni (Thorell, 1870) July 1994, lO', MT, 25 July 1994, lO', WT, 2 Au Material: NTI Mosvik: Meltingen (EIS 97), 27 gust 1996, Id', WT, 16 July 1996, 2d'd', MT, 2 August 1996,299, MT, 13 September 1997, lO', August 1996, lO',MT.All from coastal sprucefor WT, coastal spruce forest (Temmeras et al. 2000). est (Temmeras et al. 2000). A. nordmanni has not been recorded from Nor This is the first record from Norway for almost a way since Collett (1877) who found it in eastern century. The species has previously been recorded parts of South Norway and Strand (1900) who from Oslo, Drammen, Valdres, Sande and Botne collected it from southern Nordland (juveniles (Collett 1877, Strand 1900, Hauge 1989). Storm only, specimens from ZMO examined by the au (1898) reported it from Trondheim but this was thor). A. nordmanni occurs south to Uppland in referred to G. bituberculata Wa1ckenaer by Tambs Sweden (Jonsson pers. comm.), in scattered loca Lyche (1941). It ranges north to Uppland in Swe lities throughout Finland (Palmgren 1974), cen den (Jonsson pers. comm.), north to about 63° in tral and northern Europe (Heimer & Nentwig Finland (Palmgren 1974) but is apparently absent 1991). It ranges east to the Far East and west to from Great Britain and Ireland (Locket et al. 1974, North-America (Esyunin & Efimik 1996, Mikhai van Helsdingen 1996). G. omoeda is otherwise lov 1997). known from continental Europe, west and central Siberia, Shakalin and as far east as Japan (Esyunin A. nordmanni belongs to the canopy fauna (Maurer & Efimik 1996, Mikhailov 1997). & Hanggi 1990, Heimer & Nentwig 1991), prob ably being characteristic of old-growth conifer The habitat preferences are probably similar to A. ous forests. It is of special conservation concern nordmanni and A. saevus, i.e. the higher layers due to its rarity and apparent sensitivity to mod and canopy of coniferous trees (Roberts 1995). ern forestry practices (Ehnstr6m & Walden 1986). Palmgren (1974) indicated that it is very stenotope, only occuring on conifers. Even ifthe species ap pear to be rather widespread in Norway, most 154 Norw. J. Entomol. 47, 153-162.2000 records are very old and the documented sensitiv L1NYPHIIDAE ity to modem forestry practices (Petterson 1996) Caviphantes saxetarum (Hull, 1916) indicate that it is vulnerable to human influence. Material: STI Midtre Gauldal: Mo (EIS 87), 13 24 May 1994, 19, PT, sand banks ( Andersen & Singa nitidula C. L. Koch, 1844 Hanssen 1994). Material: STI Midtre Gauldal: Mo (EIS 87), 13 New to Fennoscandia. C. saxetorum is a rare spe 24 May 1994, ld', PT, open sand banks (Andersen cies known from scattered localities north to Scot & Hanssen 1994). land in Great Britain (Cooke & Merrett 1967, New to Norway. Apparently a rather rare species Locket et al. 1974), central continental Europe in both Sweden where it is known from Uppland (Georgescu 1973, Wunderlich 1975, 1979b, Mau and Dalame (Jonsson and Kronestedt pers. comm.) rer & Hanggi 1989, Steinberger 1996, Thaler and Finland where it has been found in a few places 1993), Poland (Starega 1972), Russia (Mikhailov in the southeast and along the coast ofthe Botnian 1997) and North America (Crawford 1990). This Bay (Palmgren 1974). is in other words the northernmost record in Eu rope and a rather surprising discovery. S. nitidula is a riparian species occurring in low vegetation and litter along streaming water (Palm Locket et al. (1974) and Steinberger (1996) indi gren 1974, Roberts 1995). cated that the species is restricted to open sand banks along rivers although Wunderlich (1975) GNAPHOSIDAE and Maurer & Hanggi (1989) reported C. saxeto rum & Micaria subopaca Westring, 1861 from dry, ruderal sites. Cooke Merrett (1967) found the species beneath stones and boul Material: STI Trondheim: Trondheim City (EIS ders on sand or shingle-covered river banks dur 92),28 November 1951,19, taken inside the Muse ing the fall and under smaller stones during the um ofNatural History and Archaeology (MNHA, summer. The spider occupies the small spaces leg. R. Dahlby). between the rocks and sand (Wunderlich 1979b). Previous Norwegian records are restricted to The spider itself is very small (1.5 - 1.7 mm total southeastern and southern Norway (Hauge 1989), length, Cooke & Merrett 1967) and pale in col and the present record is the northernmost from our. The vulval structures (Figure 1) ofthe present Europe. In Sweden only known as far north as specimen correspond to the photograph provided Uppland (Jonsson pers. comm., Tullgren 1946), while there are only scattered records from south ern Finland (Palmgren 1943). M. subopaca is con sidered rare in Great Britain (Leighton 1971) whe re records are concentrated around London (Locket et al. 1974), but is relatively common in continen tal Europe (Heimer & Nentwig 1991). ~) 1'~:'IJ'!v;. The species is synanhtropic in the northern end of 't,),,\jl\~1 "~\ll'-'i:~~:))0 its range and distribution data suggest a transition C _ '. '/ I I, in habitat preference at the margins of its range "'\:\' : c? both in regard to altitude and latitude. Roberts -=-=~~I~ ... \. ) .- (1995) mentions it being abundant in a Liverpool factory, all Finnish records are from inside houses ~~l ~h.. (Palmgren 1943) and according to Thaler (1997) Q.bO D it is «an der Waldgrenze synantrop». Its natural habitat is tree trunks, epecially in sunny places (Miller 1967, Wunderlich 1979a, Roberts 1995, Figure 1. Caviphantes saxetorum. Vulvae. Thaler 1997). 155 Aakra: Records of spiders (Araneae) from central Norway by Cooke & Merrett (1967). The coiled tubules seen Tube No. 506. 299, 1juv., June 1875, leg. R. Col to the side and below the spermatheca are very char lett, found in Tube No. 708 labeled < 156 Norw. J. Entomol. 47, 153-162.2000 Some doubt remains as to whether the species is Arctosa stigmosa (Thorell, 1875) present today along all rivers mentioned above. For Material: STI Midtre Gauldal: Follstad (ElS 87), instance, regulation ofrivers prevents deposition of 13 May-20 June 1994, 23d'd', 899, PI. Frosetoya silt and sand and causes the habitat ofA. cinerea to (EIS 87), 20 June-24 July 1994, Id', 19, PI. Mo be covered by vegetation. It shows considerable (ElS 87), 13 May-24 July 1994, 6d'd', 1subad., 399, sensitivity to habitat changes (Framenau 1995) and PT (all Andersen & Hanssen 1994). Melhus: Melhus has disappeared from at least one Norwegian local (ElS 92), 4 May 1994, ld', 19 (both donated to the ity; STI Melhus: Kuora, where the habitat of the Swedish Museum ofNatural History), manual col- species no longer exists (1. Andersen pers. comm.). Figure 2. Arctosa cinerea. Habitus, adult female. 157 Aakra: Records of spiders (Araneae) from central Norway lecting. Gravnik, S0berg, 22 May 2000, 19, hand developing vegetation cover and other factors re collecting from litter (KA). Orkdal: Elnvang (EIS duced the number ofavailable habitats populations 91),26 May 2000,19, manual collecting, (OF). were increasingly isolated until only those inhab iting sandy riverbanks remained. The species may New to Fennoscandia. This is the northernmost eventually be found in appropriate habitats in record from Europe and a very surprising discov Sweden (Klariilven) and Finland. Krogerus (1932) ery because it is a fairly large spider being abun failed to collect it from sandy coastal dunes in Fin dant in the appropriate habitat. Similar patterns of land however, and A. stigmosa has never been isolated occurences have been reported in ripar found in this type of habitat. The other large riv ian beetles, however (Andersen 1983, 1984, An ers in central Norway probably harbour A. stig dersen & Hanssen 1992). mosa as well, at least those where A. cinerea has The closest known occurences are from Baltikum been found (see above). (Relys 1994, Mikhailov 1997) and Poland (Kupry Ecological requirements of A. stigmosa are very janowicz pers. comm.). It is also known from cen similar to those ofA. cinerea, all specimens were tral parts ofcontinental Europe, including France, collected in vegetation-free or barely vegetated Romania, Bulgaria, Hungary, Slovakia, southern sand and gravel banks alongside the rivers Gaula Germany, Switzerland and ranges as far east as and Orkla which is in accordance with the habitat southern Siberia, China, Japan and Korea (Denis 1937, Buchar 1968, Blick & Scheidler 1991, Paik 1994, Framenau 1995, Steinberger 1996, Esyunin & Efimik 1996, Marusik et al. 1996, Mikhailov 1997, Yin et al. 1997, Song et al. 1999). In conti nental parts of Europe it is very rare and local (Buchar & Thaler 1995, Steinberger 1996). The Norwegian populations reported here are thus isolated ones, probably ofrelictual character. Pre sumably A. stigmosa colonized the region some time after the retreat of the ice cover when sandy river and glacial deposits were commonplace. As B A Figure 3. Arctosa stigmosa. Epigyne (A) and male palpal organ (8). 158 OOOG "G9t-f;9t 'Lt>' ",OW01U3 r "WON Aakra: Records of spiders (Araneae) from central Norway description of Heimer & Nentwig (1991) and gren 1943) and is otherwise widely distributed in Buchar & Thaler (1995). Marusik et al. (1996) col Europe and throughout Asia as far east as the Kuri lected the species in «stony shore in forest steppe». Ie islands (Mikhailov 1997). Available literature does not reveal much about The habitat descriptions ofthis species vary some the life history of this species. It not known if the what, Heimer & Nentwig (1991) suggest that M. mode oflife is similartoA. cinerea (see Framenau jOlmicaria may be found in a wide variety ofhabi 1995). The large number ofspecimens caught com tats. The current record is more in accordance with pared to A. cinerea may suggest a cursorial hunt Tullgren (1944) who indicated a preference for ing style, but could also be explained by the smaller spaces beneath flat rocks on warm beaches. size ofA. stigmosa. Main activity in central Nor way (based on the pitfall material) appears to take place in early summer (May-June), but specimens THERIDIIDAE of both sexes were found until late July. This cor Dipoena torva (Thorell, 1875) responds to the activity period given by Heimer & Material: NTI Mosvik: Meltingen (ElS 97), 2 Nentwig (1991), although they indicate that adults August 1996, 1cr, MT, coastal spruce forest (T0m may also be found in October. No specimens have meras et al. 2000). been seen running in daylight, indicating that the species is nocturnal. A single female has been col Northernmost record in Europe. The species was lected from litter in the forest bordering the sand recently reported from Norway for the first time banks while another was found beneath stones. It (Aakra 2000), details regarding distribution, habi is therefore likely that the species spends the day tat preferences and conservation status were given in such places and not in burrows. therein. The specimen reported here is the first male collected in Norway. The epigyne (Figure 3A) is distinctive and in good agreement with depictions given by Denis (1937), Acknowledgements. Thanks to Johan Andersen, Odd Miller (1971) and Heimer & Nentwig (1991). The var Hanssen and Frode 0degaard for help with the male palal organ is distinguished from other mem material from Gaula, to the staff at ZMO for their kind bers of the genus by the shape ofthe tegular apo assistance during my visit, to Bjorn Age Tommmerils physis which is distinctly curved and bears a small from NINA for permission to publish their material from knob and pointy projections on the posteriolateral Gaula, Mosvik and Hjerkinn, to Arne Bretten for ac side (Figure 3B). The habitus ofthe male is shown cess to his collection, to Erling Hauge and Lars Ove Hansen for comments on the manuscript, to Theo Blick in Figure 4, the female is very similar. The black for information on various species and to Torbjorn ocular region ofthe carapace contrasting with the Kronestedt for verifying the determination ofArctosa lighter, brownish areas behind is characteristic as stigmosa. Also thanks to Oddvar Hanssen for prepar is the light yellow-greenish cardiac mark clearly ing Euparal mounts of the epigynes of Caviphantes visible against the dark brown surrounding areas saxetorum and A. stigmosa and Otto Frengen for his on the abdomen. All legs are annulated in dark excellent company and help on my collection trips. brown. References SALTICIDAE Aakra, K. 2000. New records ofspiders (Araneae) from Myrmarachne formicaria (De Geer, 1778) Norway with notes on epigynal characters of Philo dromus fuscomarginatus (De Geer) (Philodromi Material: STI Midtre Gauldal: Mo (ElS 87), 13 dae) and Araneus sturmi(Hahn) (Araneidae). Norw. 24 May 1994, 1cr, PT, sandy ground in Salix 1. Entomo\. 47, 77-88. Triandra forest (Andersen & Hanssen 1994). Almquist, S. 1973. Spider associations in coastal sand dunes. Oikos 24, 444-457. New to Norway.This is the northernmost record Andersen, J. 1983. Towards an ecological explanation in Europe. The species has been taken north to of the geographical distribution of riparian beetles Varmland in Sweden (Jonsson pers. comm.), is in western Europe. J. Biogeogr. 10,421-45. known from the southern coast ofFinland (Palm Andersen, J. 1984. Gaula, et vassdrag med en unik 160 Norw. J. Entomol. 47, 153-162.2000 elvebreddfauna. Insekt-Nytt 9 (1), 21-27. Georgescu, M. 1973. Le developpement postembryon Andersen, J. & Hanssen, O. 1992. Lopebiller med isolert naire de I'especie cavernicola Lessertiella dobrogi forekomst i Norge. Tnsekt-Nytt 17, 5-16. ca Dumitrescu & Miller (Araneida, Micryphantidae). Andersen, J. & Hanssen, O. 1994. Invertebratfaunaen Trav. Inst. Speo!. «E. Racovitza» 12,63-73. pa elvebredder - ett oversett element. I. Biller (Cole Gudik-Sorensen, O. 1997. Troxochrus nasutus Schen optera) ved Gaula i Sor-Trondelag. NINA Opp kel, 1925 in Denmark (Araneae, Linyphiidae). Ent. dragsmelding 326, 1-23. Meddr. 65, 39--40. Andersen, T. & Hauge, E. 1995. Pitfall catches of spi Hauge, E. 1972. Spiders and Harvestmen from More ders (Araneae) from proposed Nature Reserves on and Romsdal and Trondelag, Norway. Norsk ent. Tjome, Vestfold, SE Norway. Fauna Norv. Ser. B Tidskr. 19, 117-12I. 42, 1-10. Hauge, E. 1989. An annotated check-list ofNorwegian Blick, T. & Scheidler, M. 1991. Kommentierte Arten spiders (Araneae). Insecta Norvegiae 4, 1--40. liste der Spinnen Bayerens (Araneae). Arachno!. Hauge, E. & Furunes, K. A. 1976. New localities for Mitt. 1,27-80. three species of spiders (Araneae) from Norway. Buchar, J.1968. Zur Lycosidenfauna Bulgariens (Ara Norw. J. Ent. 23, 87. chn., Araneae). Vest. cs. Spoelc. zoo!. 32, 116-130. Heimer, S. & Nentwig, W. 1991. Spinnen Mitteleuropas. Buchar, J. & Thaler, K. 1995. Die Wolfspinnen von Ein Bestimmungsbuch. 543 pp.Verlag Paul Parey. Osterreich 2: Gattungen Arctosa. Tricca. Trochosa Holm. A 1947. Svensk spindelfauna. 3. Egentliga (Arachnida, Lycosidae). Faunistisch-Tiergeogra spindlar. Araneae. 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Die Spinnenfauna der Ufer accepted 3 October 2000 lebensriiume des Lech (Nordtirol, Osterreich) (Arachnida: Araneae). Ber. nat.-med. Verein Inns bruck 83, 187-210. Storm, V. 1898. Iagtagelser over Arachnider i Trond hjems omegn. K. norske Vidensk. Selsk. Skr. 1898 (7), 1-10. Strand, E. 1898. Einige fundorte fUr Araneiden im siidlischen Norwegen. Verh. zool.-bot. Ges. Wien 48,401--404. Strand, E. 1900. Zur Kenntnis derArachniden Norwe gens. K. norske Vidensk. Selsk. Skr. 1900, 1--46. Strand, E. 1904a. Die Dictyniden, Dysderiden, Drassi 162 NolW. J. Entomol. 47, 163-176.2000 Ichneumonidae (Hymenoptera) new for the fauna of Norway Matthias Riedel, Lars Ove Hansen & 0istein Berg Riedel, M., Hansen, L. O. & Berg, 0. 2000. Ichneumonidae (Hymenoptera) new for the fauna of Norway. Norw. J. Entomoi. 47,163-176. In this faunistic survey we give distributional records for 169 species of parasitoid wasps of the family Ichneumonidae (Hymenoptera) which are new for the fauna ofNorway. Fifteen ofthem have not been reported from Scandinavia before. Key words: Ichneumonidae, Hymenoptera, fauna ofNorway. Dr. Matthias Riedel, Am Hamberg 8, D-29683 Fallingbostel, Germany. Lars Ove Hansen, Zoological Museum, Univ. ofOslo, Sarsgate J, NO-0562 Oslo, Norway. 0istein Berg, Kitty Kiellands vei 19 C, NO-1344 Haslum, Norway. INTRODUCTION MATERIAL With more than 21,000 valid extant species world The Ichneumonidae recorded in this paper were wide the Ichneumonidae represent one ofthe larg mostly collected in the Southern and Southeast est families ofthe Hymenoptera (Yu 1999). About ern parts of the country by the use of hand nets 4,000-4,500 species have been found in Europe and Malaise traps. To avoid severe misinterpreta so far (Gauld & Bolton 1988). Despite their abun tions, we excluded the species which have not been dance and important role in biological pest con identified with certain accuracy. trol, the taxonomy, ecology, and geographical dis The subfamilies, tribes, genera, and species are tribution of many groups of Ichneumonidae re listed in alphabetic order using the recent Interac main incompletely known, even in better studied tive Catalogue ofWorld Ichneumonidae (Yu 1999). areas such as Europe. From this catalogue we also took the distributional To establish a better knowledge of the diversity records, but mention only the Scandinavian coun and distribution ofNorwegian Ichneumonidae, we tries in more detail here. The biogeographic re studied collections of ichneumon wasps which gions within Norway are defined in accordance to have been brought together from several locali 0kland (1981 ). The reference material for our pub ties during the last years. From this material more lication is kept in the personal collection of the than 700 different ichneumonid species have been first author (M. Riedel), and at the Zoological mu identified yet. The present publication is the sec seum, Oslo. ond one covering these findings with the aim to The following abbreviations has been used in the compile a catalogue of all Ichneumonidae found text: ABa =AlfBakke, BAS = BjornA. Sagvolden, in Norway. The first was presented by Riedel & GWa = Gaute Walberg, KMy = Kai Myhr, LOH = Berg (1997). Lars Ove Hansen, MFa = Morten Falck, PAT = Per A. Tangen, no = Thor Jan Olsen, YBe = Yngvar Berg, 0Be = 0istein Berg, MT = Malaise trap, ZMO = Zoological museum, Oslo. 163 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway LIST OF SPECIES BANCHINAE ACAENITINAE Teleutaea striata (Gravenhorst, 1829) Coleocentrus excitator (Poda, 1761) TEl Notodden: Lisleherad (EIS 27), 19 6 August - 19 October 1993, MT, leg. ABa; B0 Hurum: AK B 164 Norw. J. Entomol. 47, 163-176.2000 Distribution: Holarctic, known from Finland and tic, known from Finland and Sweden. Sweden. Rhimphoctona obscuripes (Holmgren, 1860) Dusona debilis (Forster, 1868) TEl Notodden: Lisleherad (EIS 27), 299 22 June VE Vale: LanglJya (EIS 19), 198 July - 2 August - 6 August 1992, MT, leg. ABa. Distribution: 1991, MT, leg. LOH. Distribution: Known from Palaearctic, known from Finland and Sweden. Germany and Finland. COLLYRIINAE Dusona disc/usa (Forster, 1868) Collyria trichophtha/ma Thomson, 1877 ON Nord-Fron: Hesteskobakken (EIS 62),4996 Our report ofCollyria coxator (Villers) from Nor August - 19 September 1992, MT, leg. KMy & way (Riedel & Berg 1997) is a misdetermination LOH. Distribution: Palaearctic, known from Fin ofthis species, Collyria coxator has not been found land. in Norway so far. B0 RlJyken: Kinnartangen (EIS 28), 19 June 1993, MT, leg. LOH; Nedre Eiker: Dusona fa/cator (Fabricius, 1775) Ryghsetra (EIS 28), 19 June 1994, MT, leg. YBe o Rygge: Sildebauen(EIS 19), 1926 August 1995, & LOH. Distribution: Palaearctic, known from leg. 0Be. Distribution: Palaearctic, known from Sweden. Finland and Sweden. CRYPTINAE Dusona vidua (Gravenhorst, 1829) Cryptini o Halden, Enningdalen, KirkeblJen (EIS 12), 19 Agrothereutes /eucorhaeus (Donovan, 1810) 10 August 1997, leg. no. Distribution: Palaearc BV Rollag: Djupdal (EIS 27), 1928 July 1993, tic, known from Finland and Sweden. leg. BAS; VE Vale: LanglJya (EIS 19), 2(J(J June 1990, leg. LOH. Distribution: Palaearctic, known Eriborus obscuripes Horstmann, 1987 from Finland and Sweden. o Sarpsborg: Tune, Rakil (EIS 20), 19 1 August 1994, leg. no. Distribution: Central Europe, Fin Hop/ocryptus rufonigra (Desvignes, 1856) land. VE Vale: Langoya (EIS 19), 1928 May - 8 July 1991 and 699 and W 8 July - 2 August 1991, MT, Leptocampop/ex punctu/atus (Ratzeburg, 1848) leg. LOH, B0 RlJyken: Kinnartangen (EIS 28), [syn. cremastoides Holmgren] 1924 July 1993, leg. 0Be; TEl Notodden: Lisle o Sarpsborg: Tune, Rakil (EIS 20), 299 20 July herad (EIS 27), 19 6 August - 9 October 1993, 1995 and 26 August 1994, leg. no. Distribution: MT, leg. ABa. Distribution: Palaearctic, known Holarctic, known from Sweden. from Finland and Sweden. Phobocampe crassiuscu/a (Gravenhorst, 1829) Hop/ocryptus quadriguttata (Gravenhorst, 1829) o Sarpsborg: Tune, Rakil (EIS 20), 19 10 May TEl Notodden: Lisleherad (EIS 27), 1922 June 1993, leg. no; 0 Rygge: Ekeby, Gunnarsbybek 6 August 1993, MT, leg. ABa. Distribution: Palae ken (EIS 19), 19 17 June - 21 July 1992, MT, leg. arctic, known from Sweden. LOH and GWa. Distribution: Palaearctic, intro duced into Canada, known from Sweden. Cryptus minator (Gravenhorst, 1829) The identity of this species is questionable (van Rhimphoctona me/anura (Holmgren, 1860) Rossem 1969). The specimen mentioned below has o Halden, Enningdalen, Kirkeboen (EIS 12), 19 a body length of 6.5 mm and length offorewing of 10 August 1997, leg. no. Distribution: Palaearc- 5.5 mm. The propodeal spiracles are ovoid, about 165 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway 2.0 times as long as wide, the gastral tergites are Mesostenus dentifer Thomson, 1896 coriaceous. Areola with strongly convergent cubital B0 Drammen: Underlia (EIS 28), W'May 1994, cross veins, 2rm almost twice as long as 3rm. TEl MT, leg. LOR; ON Nord-Fron: Resteskobakken Notodden: Lisleherad (EIS 27), 1927 May - 22 (EIS 62), 599 and 30'0' 6 August - 19 September June 1993, MT, leg. ABa. Distribution: Palaearctic, 1992, MT, leg. KMy & LOR. Distribution: Palae known from Denmark, Finland, and Sweden. arctic, known from Sweden. Enclisis maci/enta (Gravenhorst, 1829) Trychosis neg/ecta (Tschek, 1871) o Sarpsborg: Tune, Rakil (EIS 20), 195 August TEl Notodden: Lisleherad (EIS 27), 499 27 May 1994, leg. no; 0 Rygge: Ekeby, Gunnarsbybek 22 June 1993, MI, leg. ABa. Distribution: Palae ken (EIS 19), 299 19 May- 17 June 1992, MT, leg. arctic, known from Finland and Sweden. LOR & GWa. Distribution: Palaearctic, known from Denmark, Finland, and Sweden. Trychosis tristator (Tschek, 1871) VE Vale: Lang0ya (EIS 19), W' 8 July - 2 August Enclisis vindex (Tschek, 1871) 1991, MT, leg. LOR, ON Nord-Fron: Restesko TEl Notodden: Lisleherad (EIS 27), 1927 May bakken (EIS 62), 19 6 August - 19 September 22 June 1993, MI, leg. ABa. Distribution: Palae 1992, MI, leg. KMy & LOR. Distribution: Palae arctic, known from Denmark, Finland, and Sweden. arctic, known from Denmark, Finland, and Swe den. Hidryta fusiventris (Thomson, 1873) B0 Reyken: Kinnartangen (EIS 28), 19 6 July - 4 Xy/ophrurus /ancifer (Gravenhorst, 1829) August 1991, leg. LOR; 0 Sarpsborg: Tune, Riikil TEl Notodden: Lisleherad (EIS 27), 1927 May (EIS 20), W' 25 July 1993, leg. no; VE Vale: 22 June 1993 and 299 22 June - 6 August 1993, Lang0ya (EIS 19),299 and 10' 8 July - 2 August MI, leg. ABa; B0 Nedre Eiker: Mj0ndalen, Mile 1991, MT, leg. LOR. Distribution: Palaearctic, tjem (EIS 28), 19 and 20'0' May 1994, MI, leg. known from Sweden. YBe & LOR; Drammen: Underlia (EIS 28), I 9 May 1994, MT, leg. LOR; Reyken: Kinnartangen /diolispa suba/pina (Schmiedeknecht, 1904) (EIS 28), 2928 May - 6 July 1991, MT, leg. LOR. FV Mas0Y: RolvS0y, Gunnames (EIS 186), 19 MI, Distribution: Palaearctic, known from Finland and August 1992, leg. PAT. Distribution: Palaearctic, Sweden. known from Finland and Sweden. Hemigasterini Listrognathus compressicornis (Gravenhorst, Aptesis cretata (Gravenhorst, 1829) 1829) o Rygge: Ekeby, Telemarkslunden (EIS 19), W' BV Rollag: Varviken (EIS 35), 19 July 1992, MI, 21 July - 24 August 1992, MI, leg. LOR & GWa. leg. BAS. Distribution: Palaearctic, known from Distribution: Palaearctic, known from Finland and Finland and Sweden. Sweden. Meringopus nigerrimus murorum (Tschek, 1872) Aptesis jejunator (Gravenhorst, 1807) TEl Notodden: Lisleherad (EIS 27), 1927 May o Rygge: Ekeby, Ielemarkslunden (EIS 19), W' 22 June 1993, MT, leg. ABa; 0 Fredrikstad: Ons 19 May - 17 June 1992, MI, leg. LOR & GWa. 0Y, Rank0, Bloksberg (EIS 20), 10' 3-29 June 1995, Distribution: Palaearctic, known from Denmark, leg. LOR & J.I.I. Batvik. Distribution: Rolarctic, Finland, and Sweden. known from Finland and Sweden. 166 Norw. J. Entomo/. 47, 163-176. 2000 Colocnema rufina (Gravenhorst, 1829) from Denmark, Finland, and Sweden. B0 Nedre Eiker: Mjondalen, Miletjem (EIS 28), 2(Je) July 1988, leg. D. Ruud. Distribution: Palae Mastrus longicauda Horstmann, 1990 arctic, known from Denmark and Sweden. VE Stokke: MeIsomvik (EIS 19), 19 22 July -27 August 1995, window trap, leg. O. Hanssen & Demopheles corruptor (Taschenberg, 1865) LOH. Distribution: Known from Denmark and B0 Humm: Tofteholmen (EIS 19), 19 7-31 July Sweden. 1991, MT, leg. LOH. Distribution: Holarctic, new record for Scandinavia. Orthizema obscurum Horstmann, 1993 BV Rollag: Viirviken (EIS 35),19 July 1992, MT, Plectocryptus digitatus (Gmelin, 1790) leg. BAS. Distribution: England and Austria, new o Rygge: Ekeby, Telemarkslunden (EIS 19), XK) record for Scandinavia 19 May - 17 June 1992, MT, leg. LOH & GWa. Distribution: Palaearctic, known from Denmark Phygadeuon elegans (Forster, 1850) and Sweden. The specimens are brachypterous. 0 Moss: Jeloy, Hvittingbukta (EIS 19), 299 30 June - 31 July Phygadeuontini 1995, pitfall-trap, leg. O. Hanssen & G. Hardeng. Bathythrix linearis (Gravenhorst, 1829) Distribution: Palaearctic, known from Denmark, Finland, and Sweden. o Sarpsborg: Riikil (EIS 20),197 July 1997, leg. no. Distribution: Palaearctic, known from Swe den. CTENOPELMATINAE Ctenopelmatini Bathythrix prominens (Strobl, 1901) Xenoschesis nigricoxa Strobl, 1903 o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, HOI Ulvik: Finse (EIS 42),1919 July 1991, leg. 1924August -16 October 1992, leg. LOH& GWa. O.J. Lonnve. Distribution: Palaearctic, new record Distribution: Palaearctic, known from Sweden. for Scandinavia. Chirotica maculipennis (Gravenhorst, 1829) Pionini TEl Notodden: Lisleherad (EIS 27) , 1922 June Pion crassipes (Holmgren, 1857) 6 August 1993, MT, leg. ABa. Distribution: Palae o Fredrikstad: Gansrod!0ra (EIS 20), 19 12 July arctic, known from Finland and Sweden. 1996, leg. no. Distribution: Palaearctic, known from Finland and Sweden. Endasys alutaceus (Habermehl, 1912) o Sarpsborg, Riikil (EIS 20), 1(J 15 June 1997, Rhorus chrysopus (Gmelin, 1790) leg. no. Distribution: Palaearctic, known from o Sarpsborg: Maugesten (EIS 20), 19 29 June Denmark, Finland, and Sweden. 1992, leg. no; B0 Nedre Eiker: Mjondalen, Miletjem (EIS 28), W July 1988, leg. D. Ruud. Endasys striatus (Kiss, 1924) Distribution: Palaearctic, known from Finland. o Sarpsborg: Tune, Riikil (EIS 20), W 10 July 1994, leg. no. Distribution: Palaearctic, known Sympherta ullrichi (Tschek, 1869) from Denmark, Finland, and Sweden. o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, 3(J(J 19 May - 17 June 1992, leg. LOH & GWa. Glyphicnemis vagabunda (Gravenhorst, 1829) Distribution: Palaearctic, known from Finland. B0 Royken: Kinnartangen (EIS 28), W 8 July 1989, leg. LOH. Distribution: Palaearctic, known 167 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway Trematopygus dubitor Hinz, 1982 Campodorus caligatus (Gravenhorst, 1829) BV Rollag: Kjomme (EIS 27), 19 13 June 1995, e Sarpsborg: Tune, Riikil (EIS 20), 1927 Octo leg. BAS. Distribution: Palaearctic, known from ber 1985, leg. no. Distribution: Palaearctic, Sweden. known from Finland and Sweden. Trematopygus lethierryi (Thomson, 1893) Campodorus clypealis (Thomson, 1893) Be Nedre Eiker: Ryghsetra (EIS 28), ld' May BVRollag: Veggli (EIS 35),1919 June 1995, leg. 1994, MT, leg. YBe & LOH. Distribution: Palae BAS. Distribution: Palaearctic, new record for arctic, known from Sweden. Scandinavia. Perilissini Hyperbatus sternoxanthus (Gravenhorst, 1829) Lathrolestes verticalis (Brischke, 1871) VE Viile: Lang0ya (EIS 19), 19 September - Oc e Rygge, Ekeby, Gunnarsbybekken (EIS 19), 1(J tober 1987, leg. LOH. Distribution: Palaearctic, 21 July - 24 August 1992, 1(J 24 August - 16 Octo known from Finland and Sweden. ber 1992, leg. LOH & GWa. Distribution: Palaearctic, known from Finland and Sweden. Lamachus coalitorius (Thunberg, 1822) VE Viile: Lang0ya (EIS 19), 198 July 1991, leg. Perilissus limitaris (Gravenhorst, 1829) LOR. Distribution: Palaearctic, introduced into e Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, Canada and U.S.A., known from Finland and 399 19 May - 17 June 1992, leg. LOH & GWa; Sweden. Be Drammen: Underlia (EIS 28), 1(J 1 October 1985, leg. LOH; Nedre Eiker: Mjemdalen, Mile Lamachus frutetorum (Hartig, 1838) tjem (EIS 28), 1(J 15 September 1988, leg. D.W.B. VE Viile: Lang0ya (EIS 19), 19 8 July - 2 August Johansen, Distribution: Palaearctic, known from 1991, MT, leg. LOH; Be Dramrnen: Underlia(EIS Denmark and Finland. 28), 19 I October 1985, leg. LOR. Distribution: Palaearctic, known from Finland and Sweden. Perilissus lutescens Holmgren, 1857 RY Randaberg: Gmdem (EIS 7), 193 June 1995, Protarchus testatorius (Thunberg, 1822) leg. 0Be. Distribution: Palaearctic, known from STI Midtre Gauldal: Elgsh0gda 700m, 19 leg. 13 Finland and Sweden. March 1996, hatched 27 April 1996 from pupa of Cimbicidae spec. attached to twig ofBetula pube Mesoleiini scens, leg. 0Be. Distribution: Palaearctic, known Alexetercoxalis (Brischke, 1871) from Finland and Sweden. e Halden: Enningdalen, Kirkeb0en (EIS 12), 19 1August 1997, leg. no, Rygge: Sildebauen (EIS Scopesis gesticulator (Thunberg, 1822) 19), 1926 August 1995, leg. 0Be, Rcide: Tomb [syn. nigricollis Gravenhorst] (EIS 20), 19 II August 1995, leg. LOR. Distribu e Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, tion: Palaearctic, new record for Scandinavia. ld' 21 July - 24 August 1992, leg. LOH & GWa. Distribution: Holarctic, known from Finland and Barytarbes adpropinquator (Gravenhorst, 1829) Sweden. Be Lier: Garsj0 (EIS 28), 1(J 4 July 1993, leg. 0Be. Distribution: Palaearctic, known from Fin land and Sweden. 168 Norw. J. Entomol. 47,163-176.2000 Euryproctini Heterischnus truncator (Fabricius, 1798) Hadrodacty/us flavofacialis (Horstmann, 2000) o Fredrikstad: Gansrod (ElS 20), 19 10 June 1996, [syn. flavifrontator Thunberg] leg. no. Distribution: Palaearctic, known from Finland. B0 Royken: Kinnartangen (ElS 28), 1cJ 29 May 1989, leg. LOH. Distribution: Palaearctic, known from Finland and Sweden. Tycherus cepha/otes (Wesmael, 1845) RY Sandnes: Dale (ElS 7), 19 21 May 1988, ICHNEUMONINAE hatched from Pammene regiana (Lepidoptera), leg. LOH. Distribution: Palaearctic, known from Alomyini Finland and Sweden. Aethecerus disc%r Wesmael, 1845 o Rygge: Ekeby, Gunnarsbybekken (ElS 19), lcJ Platylabini 21 July - 24 August 1992, MT, leg. LOH & GWa. P/aty/abus decipiens Wesmael, 1848 Distribution: Palaearctic, known from Finland and Sweden. B0 Nedre Eiker: Mjondalen, Miletjem (ElS 28), light trap, 1cJ June 1988, leg. LOH. Distribution: Palaearctic, known from Finland and Sweden. Centeterus majorWesmael, 1845 VE Vale: Langoya (ElS 19), 1cJ 8 July - 2 August Heresiarchini 1991, MT, leg. LOH; 0 Sarpsborg: Rakil (ElS 20), 19 15 July 1995, leg. no. Distribution: Palae Amb/yjoppa fuscipennis (Wesmael, 1845) arctic, known from Finland and Sweden. B0 Humm: Molen (ElS 19), W 1 August 1991, leg. LOH; Royken: Kinnartangen (ElS 28), 19 Dicae/otus punctiventris (Thomson, 1891) June 1993, MT, leg. LOH; Nedre Eiker: Ryghsetra (ElS 28), W June 1994, at light, leg. D. Ruud. AK Oslo: Kvremer (ElS 28), 19 12-26 June 1990, VE Sande: Kommersoya (ElS 19), 2cJcJ 28 May leg. MFa. Distribution: Palaearctic, known from 9 July 1991, leg. LOH.; BV RoIJag: Rollag (ElS Sweden. 35), 19 18 June 1993, leg. BAS. Distribution: Pa laearctic, known from Finland. Erip/atys ardeicollis (Wesmael, 1845) o Sarpsborg: Skjeberg, Grimsoy (ElS 20), 1922 Amb/yjoppa proteus (Christ, 1791) April 1992, leg. B0 Drammen: Underlia no; AK Oslo: Noklevann (ElS 28), 19 11 July 1992, (ElS 28), 19 June 1993, MT, leg. LOH. Distribu leg. MFa. Distribution: Palaearctic, known from tion: Palaearctic, known from Finland and Swe Finland and Sweden. den. Coelichneumon opu/entus (Taschenberg, 1871) Heterischnus anoma/us (Wesmael, 1857) B0 Humm: Tofteholmen (ElS 19), 19 1 Septem This species is probably a colour fonn ofHeteris ber - 26 October 1991, MI, leg. LOH. Distribu chnus truncator (Fabricius). B0 Nedre Eiker: tion: Palaearctic, known from Finland and Swe Mjondalen, Miletjem (ElS 28), W 13 July 1988, den. leg. D.W.B. Johansen; 0 Rygge: Ekeby, Gunnars bybekken (ElS 19), W 17 June - 21 July 1992, MT, leg. LOH & GWa. Distribution: Palaearctic, Coe/ichneumon orbitator (Thunberg, 1822) known from Finland. o Sarpsborg: Rakil (ElS 20), 1cJ 4 June 1996, leg. no. Distribution: Palaearctic, known from Swe den. 169 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway Coelichneumon pumilionobilis Heinrich, 1951 Homotherus varipes (Gravenhorst, 1829) TEl Notodden: Lisleherad (EIS 27), 1922 June e Sarpsborg: Skjebergdalen (EIS 20), 19 7 July 6 August 1993, MT, leg. ABa. Distribution: Rol 1996, leg. no. Distribution: Palaearctic, known arctic, new for Scandinavia. from Denmark, Finland, and Sweden. Ichneumonini Ichneumon analis Gravenhorst, 1829 Cratichneumon coruscator (Linnaeus, 1758) e Sarpsborg: Tune, Rakil (EIS 20), W 25 Au VE Larvik: Brunlanes, Lerungsand (EIS 11), 2aa gust 1993, leg. no. Distribution: Palaearctic, new 23 June 1996, leg. 0Be; TRY Tromso: Telegraf for Scandinavia (a questionable record was given bukta, W 21 July 1994, leg. H. Elven. Distribu by Strand, 1913 from YAY). tion: Palaearctic, new for Scandinavia. Ichneumon minutorius Desvignes, 1856 Cratichneumon infidus (Wesmael, 1848) VE Larvik: Brunlanes, Lerungsand (EIS 11), 19 e Rygge: Ekeby, Gunnarsbybekken (EIS 19), 19 6 July 1996, leg. 0Be. Distribution: Palaearctic, 21 July- 24 August 1992, leg. LOR & GWa. Dis known from Finland and Sweden. tribution: Palaearctic, known from Finland and Sweden. Ichneumon tuberculipes Wesmael, 1848 . e Aremark: Boens::etre (EIS 21), 195 June 1994, Diphyus bicingulatus (Gravenhorst, 1829) 1922 June 1996, leg. no; Rvaler: Kj::erkoy, 19 BV Rollag: Rollag (EIS 35), 9 18 June 1993, leg. 5 July 1994, leg. LOR. VE Larvik: Brunlanes, BAS; Rollag: St::emes (EIS 27), 9 May 1994 and Lerungsand (EIS 11), 1923 June 1996, leg. 0Be; 9 20 April 1994, leg. BAS. Distribution: Palae TEl Notodden: Lisleherad (EIS 27), 19 27 May arctic, known from Finland and Sweden. 22 June 1993, MT, leg. ABa. Distribution: Palae arctic, known from Finland and Sweden. Diphyus castanopyga (Stephens, 1835) Patrocloides lapponicus (Holmgren, 1871) YAY Flekkefjord: Ridra, Dragoy (EIS 4), 19 6 May - 20 June 1982 MT, leg. A.J. Nielsen. Distri HOI Ulvik: Finse (EIS 42), 399 22 and 26 July bution: Palaearctic, known from Sweden. 1991, leg. OJ. Lonnve. Distribution: Palaearctic, known from Finland and Sweden. Diphyus indocilis (Wesmael, 1845) Stenobarichneumon basalis Perkins, 1960 e Sarpsborg: Skjeberg, Skjebergdalen (EIS 20), [syn. basiglyptus auct. nec Kriechbaumer] 19 24 April 1993, leg. no. Distribution: Palae arctic, known from Finland and Sweden. e Sarpsborg: Tune, Rakil (EIS 20), W 15 June 1995, leg. no. Distribution: Palaearctic, known from Finland. Diphyus palliatorius (Gravenhorst, 1829) Be Rurum: Skatvedt (EIS 28), W 18 July 1994, Stenobarichneumon citator (Thunberg, 1822) leg. 1. Engdal. Distribution: Palaearctic, known from Finland and Sweden. e Sarpsborg: Tune, Rakil (EIS 20) (EIS 20), 19 21 June 1994, leg. no. Distribution: Palaearctic, known from Finland and Sweden. Diphyus trifasciatus (Gravenhorst, 1829) e Sarpsborg: Tune, Rakil (EIS 20), 2<)9 10 and Thyrateles haereticus (Wesmael, 1854) 15 April 1994, leg. no; AK Asker: Rval (EIS 28), 19 30 April 1995, leg. 0Be. Distribution: e Fredrikstad: Fredrikstad (EIS 20), 19 1August Palaearctic, new for Scandinavia. 1991, leg. O. Sorlibraten; AKB::erum: Kjaglidalen (EIS 28), 19 11 - 22 June 1990, MT, leg. MFa; 170 Narw. J. Entamal. 47, 163-176.2000 Asker: Nesoya (EIS 28),899 hatched from pupa Astiphromma tenuicorne Thomson, 1886 of Inachis io (Linnaeus) (Lepidoptera) in July BO Royken: Kinnartangen (EIS 28), 196 July 1993 leg. LOH; HEN Rendalen: Ytre Rendal, 1989, leg. LOH; Humm: Tofteholmen (EIS 19), Storsjoen (EIS 64),1927 July 1945, leg. L.R. Nat 197-31 July 1991, MT, leg. LOH. Distribution: vig col. ZMO. Distribution: Palaearctic, known Palaearctic, known from Finland and Sweden. from Finland and Sweden. According to Hinz (1973) this species is mainly a parasitoid ofAglais Mesochorus brevipetiolatus Ratzeburg, 1844 urticae (Linnaeus); I. io seems to be an exceptio nal host at least in Central Europe. BO Nedre Eiker: Mjondalen, Miletjern (EIS 28), 299 June 1988, light trap, leg. LOH. Distribution: Palaearctic, known from Finland and Sweden. Virgichneumon maculicauda (Perkins, 1953) o Fredrikstad: Onsoy, Rauer (EIS 19), 1r:J 29 July Mesochorus pictilis Holmgren, 1860 1989, leg. LOH. Distribution: Palaearctic, new for Scandinavia o Rygge: Ekeby, Gunnarsbybekken (ElS 19),399 21 July - 24 August 1992, MT, leg. LOH & GWa. Distribution: Holarctic, known from Finland and Vulgichneumon deceptor (Scopoli, 1763) Sweden. ON Nord-Fron: Hesteskobakken (EIS 62), 1r:J II July 1994, leg. 0Be. Distribution: Palaearctic, Mesochorus testaceus Gravenhorst, 1829 known from Finland and Sweden. BO Royken: Kinnartangen (EIS 28), 196 July 1989, MT, leg. LOH. Distribution: Palaearctic, Vulgichneumon saturatorius (Linnaeus, 1758) known from Finland and Sweden. BO Royken: Kinnartangen (EIS 28), W 17 July 1989, leg. LOH. Distribution: Palaearctic and Ori METOPIINAE ental, known from Denmark, Finland, and Swe den. Exochus albicinctus Holmgren, 1873 var. Temples enlarged, costulae absent, head, thorax Zanthojoppa /utea (Gravenhorst, 1829) and gaster black, legs with coxae predominantly black, all femora apically, basal ring and dorsal AK B Astiphromma dorsaIe (Holmgren, 1860) Exochus ratzeburgi Holmgren, 1858 o Sarpsborg: Tune, Rflkil (ElS 20), Ir:J 15 June BO Drammen: Underlia (EIS 28), 1r:J July 1994, 1994, leg. no; Fredrikstad: Fredrikstad (EIS 20), MT, leg. LOR. Distribution: Northern and East 19 1 August 1991, leg. O. Sorlibrflten. Distribu ern Europe, known from Finland and Sweden. tion: Palaearctic and Oriental, known from Fin land and Sweden. 171 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway Metopius brevispina Thomson, 1887 Megastylus flavopictus (Gravenhorst, 1829) TEl Seljord: Blika (EIS 26), IcJ 5 - 8 August 1993, VE Ville: Lang0ya (EIS 19), 1cJ 8 July - 2 August leg. H. Elven & A. Dahl. Distribution: Palaearctic, 1991, MI, leg. LOH. Distribution: Holarctic, known known from Finland. from Denmark, Finland, and Sweden. Triclistus aethiops (Gravenhorst, 1829) Megastylus impressor Schil2ldte, 1838 VE Sande: Kommerseya (EIS 19), 1928 May - 9 FVMasey: Rolvs0Y, Gunnarnes (EIS 186), 19MI July 1991, leg. LOH. Distribution: Palaearctic, August 1992, leg. PAT. Distribution: Ho1arctic, known from Finland and Sweden. known from Denmark, Finland, and Sweden. Trieces tricarinatus (Holmgren, 1858) Megastylus orbitator Schil2ldte, 1838 o Sarpsborg: Sandbakken (EIS 20), IcJ 10 July TEl Notodden: Lisleherad (EIS 27), 196 August 1996, leg. no; VE Vale: Langeya (ElS 19), 199 - 19 October 1993, MI, leg. ABa; 0 Rygge: Eke 8 July - 2 August 1991, MI, leg. LOH. Distribu by, Gunnarsbybekken (EIS 19), MI, 299 21 July tion: Pa1aearctic, known from Finland and Swe - 24 August, leg. LOH & GWa; Rygge: Ekeby, den. Ie1emarkslunden (EIS 19), 19 17 June - 21 July 1992 and 19 21 July - 24 August 1992, MI, leg. ORTHOCENTRINAE LOH & GWa; AK Serum: L0renfallet, Egner (EIS Aperileptus microspilus Forster, 1871 37), 19 October 1994, MI, leg. LOH & O. S0rli brilten; ON Nord-Fron: Hesteskobakken (EIS 62), o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MI, 19 6 August - 19 September 1992, MI, leg. KMy 19 24 August - 16 October 1992, leg. LOH & & LOH. Distribution: Holarctic, known from Fin GWa. Distribution: Germany, Finland, and Swe land. den Orthocentrus asper Gravenhorst, 1829 Gnathochorisis crassulus (Thomson, 1888) AK Brerum: Kjaglida1en (EIS 28), 1926 May o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MI, 11 June 1990, MI, leg. MFa; 0 Rygge: Ekeby, 299 24 August - 16 October 1992, leg. LOH & Gunnarsbybekken (EIS 19), 19 24 August - 16 GWa. Distribution: Holarctic, known from Fin October 1992, MT, leg. LOH & GWa. Distribu land and Sweden. tion: Holarctic, known from Finland and Sweden. Helictes erythrostoma (Gmelin, 1790) Orthocentrus sannio Holmgren, 1858 BO Hurum: Ramvikho1men (EIS 19), 1cJ 12 Au o Rygge: Ekeby, Gunnarsbybekken (EIS 19), 1cJ gust 1990, leg. LOH. Probably 19 belonging to 24 August - 16 October 1992, MI, leg. LOH & this species: TEl Notodden: Lisleherad (EIS 27), GWa. Distribution: Palaearctic, known from Fin 96August - 19 October 1993, MI, leg. ABa. Dis land and Sweden. tribution: Holarctic, known from Finland and Sweden. Pantisarthrus luridus Forster, 1871 TEl Notodden: Lisleherad (EIS 27), 196 August Megastylus excubitor (Forster, 1871) - 19 October 1993, MT, leg. ABa; AK S0rum: HEN Rendalen: Ytre Rendal, Solbakken (EIS 64), Lerenfallet, Egner (EIS 37), 1cJ October 1994, 1925 August 1957, leg. L.R. Natvig, col. ZMO. MI, leg. LOH and S0rlibraten; 0 Sarpsborg: Distribution: Palaearctic, known from Sweden. Tune, Rakil (EIS 20), 1cJ 22 May 1994, leg. no. Distribution: Holarctic, known from Greenland, Finland and Sweden. 172 Norw. J. Entornol. 47, 163-176.2000 Proclitus praetor (Haliday, 1871) Phrudus monilicornis Bridgman, 1886 o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, B0 Drammen: Underlia (EIS 28),19 June 1992, ld' 21 July - 24 August 1992, leg. LOH & GWa; MT, leg. LOH. Distribution: Palaearctic and Ori B0 Drammen: Underlia (EIS 28), 19 October ental, known from Finland and Sweden. 1994, MT, leg. LOH. Distribution: Ho1arctic, known from Denmark, Finland, and Sweden. PIMPLINAE Ephialtini ORTHOPELMATINAE Dolichomitus pterelas (Say, 1829) Orthopelma mediator (Thunberg, 1822) o Sarpsborg: Tune, Rakil (EIS 20), 197 August B0Nedre Eiker: Ryghsetra (EIS 28),19 and 3d'd' 1995, leg. no. Distribution: Holarctic, new for May 1994, hatched from galls on Rosa sp., leg. Scandinavia. LOH; AK Ra::lingen: Tveter, 1699 and 12d'd' May 1994, hatched from galls on Rosa sp., leg. LOH. Ephialtes brevis Morley, 1914 Distribution: Holarctic, known from Finland and Sweden. BV Rollag: Bratasen (EIS 35), 19 July 1994, MT, leg. LOH and BAS; Rollag: Varviken (EIS 35), 19 August 1992, MT, leg. BAS. AK Fet: Nordre OXYTORINAE 0yeren, Ekholt (EIS 29), 192 May 1988, leg. K. Oxytorus armatus Thomson, 1883 Sund, col. ZMO; TEl Notodden: Lisleherad (EIS B0 Nedre-Eiker: Mjondalen, Miletjem (EIS 28), 27), 799 22 June - 6 August 1993 and 196 Au ld' September 1988, leg. D.W.B. Johansen. Dis gust - 19 October 1994, MT, leg. ABa. Distribu tribution: Palaearctic, known from Finland and tion: Holarctic, known from Finland. Sweden. Fredegunda dilutus (Ratzeburg, 1852) PAXYLOMMATINAE o Fredrikstad: Fredrikstad (EIS 20), 19 I August Hybrizon buccatum (Brebisson, 1825) 1991, leg. O. Sorlibraten. Distribution: Palaearctic, o Sarpsborg: Tune, Rakil (EIS 20), 499 16-20 known from Finland and Sweden. July 1994, 8August 1995,20 July 1995, leg. no; B0 Royken: Kinnartangen (EIS 28), 1928 May Liotryphon caudatus (Ratzeburg, 1848) 6 July 1991, leg. LOH; Drammen: Underlia (EIS TEl Notodden Lisleherad (EIS 27), 1927 May 28), 299 June 1992, MT, leg. LOH; VE Vale: 22 June 1993, MT, leg. ABa. Distribution: Palae Langoya (EIS 19), 299 8 July - 2 August 1991, arctic, introduced into U.S.A. and New Zealand, leg. LOH; Stokke: Melsomvik (EIS 19), 196 June known from Sweden. 22 July 1995, pitfall trap, leg. O. Hanssen & LOH; TEY Porsgrunn: Helleasen (EIS 11), 19 13 July Scambus pomorum (Ratzeburg, 1848) 27 August 1995, pitfall-trap, leg. O. Hanssen & LOH. Distribution: Palaearctic, known from Fin A variant specimen with red marks on thorax. B0 land. Nedre Eiker: Ryghsetra (EIS 28), 19 May 1994, MT, leg. YBe and LOH. Distribution: Palaearctic, known from Sweden. PHRUDINAE Astrenis paradoxa Schmiedeknecht, 1907 Tromatobia ornata (Gravenhorst, 1829) o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, o Sarpsborg: Rakil (EIS 20), Tune, 19 20 Sep 19 21 July - 24 August 1992, leg. LOH & GWa. tember 1994, leg. no. Distribution: Palaearctic, Distribution: Central Europe, new for Scandina known from Finland and Sweden. via. 173 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway Zaglyptus mu/ticolor (Gravenhorst, 1829) RHYSSII\IAE TEl Notodden: Lisleherad (EIS 27), 196 August Megarhyssa perlata (Christ, 1791) - 19 October 1993, MT, leg. ABa; BO Royken: TEl Notodden: Lisleherad (EIS 27), 196 August Kinnartangen (EIS 28), I(J 8 September - 24 Oc - 19 October 1993, MT, leg. ABa. Distribution: tober 1991, MT, leg. LOR. Distribution: Palaearc Palaearctic, known from Sweden. tic and Oriental, known from Finland and Swe den. TERSILOCHINAE Barycnemis gracil/ima Thomson, 1889 Polysphinctini BV Rollag: Rollag (EIS 35), 195 June 1995, leg. Zatypota albicoxa (Walker, 1874) BAS. Distribution: Palaearctic, known from Fin BO Rumm: Molen (EIS 19), 192 - 4 July 1990, land and Sweden. leg. LOR; Drammen: Underlia (EIS 28), 19 July 1992, MT, leg. LOR. Distribution: Palaearctic and Phradis interstitialis (Thomson, 1889) Oriental, known from Denmark and Sweden. ON Nord-Fron: Resteskobakken (EIS 62), 399 1-25 May 1992, MT, leg. KMy & LOR. Distribu Pimplini tion: Palaearctic, known from Denmark, Finland, Apechthis capulifera (Kriechbaumer, 1887) and Sweden. o Aremark: Boensletre (EIS 21), 195 June 1994, leg. no. Distribution: Palaearctic, known from Probles microcephalus (Gravenhorst, 1829) Sweden. BO Royken: Kinnartangen (EIS 28), 19 11 July 1991, leg. LOR. Distribution: Palaearctic, known POEMENIINAE from Denmark, Finland, and Sweden. Deuteroxorides elevator (Panzer, 1799) TEl Tinn: Riikenes (EIS 26), J(J July 1995, MT, Spinolochus laevifrons (Holmgren, 1860) leg. BAS. Distribution: Palaearctic, known from o Sarpsborg: Tune, Riikil (EIS 20), 19 30 July Finland and Sweden. 1994, leg. TIO. Distribution: Palaearctic, known from Finland and Sweden. Poemenia brachyura Holmgren, 1860 o Sarpsborg: Tune, Riikil (EIS 20), 299 10 June TRYPHONINAE 1993, IOAugust 1994, leg. TIO. Distribution: Pal Tribus Phytodietini aearctic, known from Finland and Sweden. Netelia tarsata (Brischke, 1880) o Sarpsborg: Riikil (EIS 20), Tune, 19 1994 Poemenia col/aris (Haupt, 1917) (without date), leg. TIO. Distribution: Rolarctic, o Sarpsborg, Sandbakken (EIS 20), 19 27 July known from Finland. 1996, leg. TIO. Distribution: Palaearctic, known from Sweden. Phyfodietus polyzonias (Forster, 1771) o Rygge: Ekeby, Gunnarsbybekken (EIS 19), MT, Poemenia notata Holmgren, 1859 19 19 May - 17 June 1992, leg. LOR & GWa. TEl Notodden: Lisleherad (EIS 27), 299 22 June Distribution: Palaearctic, known from Denmark, - 6 August 1993, MT, leg. ABa. Distribution: Pal Finland, and Sweden. aearctic, known from Finland and Sweden. 174 Norw. J. Entomol. 47, 163-176.2000 Tryphonini Eridolius schiodtei (Holmgren, 1857) Cosmoconus nigriventris Kasparyan, 1971 () Rygge: Ekeby, Gunnarsbybekken (EIS 19),200 B(} Nedre Eiker: Mjondalen, Miletjem (EIS 28), 17 June - 21 July 1992 and 2099 24 August - 16 299 and 200 31 August 1988, leg. D.W.B. Johan October 1992, MT, leg. LOH & GWa. Distribu sen. Distribution: Palaearctic, known from Fin tion: Known from Finland, Russia, and Sweden. land. Excavarus apiarius (Gravenhorst, 1829) Ctenochira angulata (Thomson, 1883) BV Rollag: Veggli (EIS 35), 199 July 1995, leg. () Sarpsborg: Sandbakken (EIS 20), W 20 June BAS; AK Sorum: Lorenfallet, Egner (EIS 37), 1a 1996, leg. no. Distribution: Palaearctic, known 7 July 1994, leg. O. Sorlibriiten. Distribution: Pal from Finland and Sweden. aearctic, known from Denmark, Finland, and Swe den. Ctenochira propinqua (Gravenhorst, 1829) Exyston pratorum (Woldstedt, 1874) BV Rollag: Kjomme (EIS 27), 19 13 June 1995, leg. BAS. Distribution: Palaearctic, known from () Sarpsborg: Sandbakken (EIS 20), 19 6 July Denmark, Finland, and Sweden. 1996, leg. no; Sarpsborg: Tune, Rilkil (EIS 20), 19 22 June 1994, leg. no; BV Rollag: Veggli (EIS 35), W 23 June 1995, leg. BAS; VE Larvik: Erromenus analis (Brischke, 1871) Brunlanes, Lerungsand (EIS II), 19 15 June 1997, () Sarpsborg: Rilkil (EIS 20), Tune, 19 22 July leg. 0Be. Distribution: Palaearctic, known from 1994, leg. no. Distribution: Holarctic, known Finland and Sweden. from Finland and Sweden. Smicroplectrus erosus (Holmgren, 1857) Neleges proditor (Gravenhorst, 1829) BV Rollag: Veggli (EIS 35), W 23 June 1995, AK Oslo: Roa (EIS 28), 197 July 1995, leg. 0Be. leg. BAS. Distribution: Palaearctic, known from Distribution: Palaearctic, new record for Scandi Sweden. navia Idiogrammatini Polyblastus pal/icoxa Thomson, 1888 Idiogramma alysiina (Thomson, 1888) AK Oslo: Kvcemer(EIS 28),1912- 26 June 1990, () Rilde: Tomb (EIS 20), 19 and 200 18 May 1995, MT, leg. MFa. Distribution: Palaearctic, known leg. Distribution: Known from Finland and from Finland and Sweden. no. Germany. Tryphon heliophilus (Gravenhorst, 1829) XORIDINAE VE Ville: Langoya (EIS 19), 198 July - 2 August Ischnoceros caligatus (Gravenhorst, 1829) 1991, MT, leg. LOH; () Sarpsborg: Tune, Rilkil (EIS 20), 19 28 July 1994, leg. no. Distribu () Rygge: Ekeby, Telemarkslunden (EIS 19), MT, tion: Palaearctic, known from Denmark, Finland, 1917 June - 21 July 1992, 19 17 June - 21 July and Sweden. 1992, leg. LOH & GWa. Distribution: Palaearctic, known from Finland and Sweden. Exenterini Xorides gravenhorstii (Curtis, 1831) Cycasis rubiginosa (Gravenhorst, 1829) BV Rollag: Djupdal (EIS 27),1928 July 1993, leg. () Sarpsborg: Rilkil (EIS 20), 299 10 July 1996 BAS; () Rygge: Telemarkslunden (EIS 19), 1921 and 20 June 1997, leg. no. Distribution: Palae July - 24 August 1992, MI, leg. LOH & GWa. Dis arctic, known from Finland and Sweden. tribution: Palaearctic, known from Sweden. 175 Riedel, Hansen & Berg: Ichneumonidae (Hymenoptera) new for the fauna of Norway Xorides irrigator (Fabricius, 1793) biogeografiske oppgaver - et revidert Strand-sys tem. Fauna (Oslo) 34, 167-178. TEl Notodden: Lisleherad (EIS 27), 399 and ld' Riedel, M. & Berg, 0. 1997. Faunistic remarks on Nor 27 May - 19 October 1993, MT, leg. ABa. Distri wegian Ichneumonidae (Hymenoptera). Fauna bution: Palaearctic, known from Finland and Swe norv. Ser. B 44, 39-53. den. Rossem, G. van. 1969. A revision ofthe genus Cryptus Fabricius S.str. in the Western Palaearctic region, Xorides niger Pfeffer, 1913 with keys to genera ofCryptina and species ofCryp tus. Tijdschr. Ent. 112,299-375. () Sarpsborg: Tune, Rakil (EIS 20),19 22 August Yu, D.S. 1999. Interactive Catalogue ofWorld Ichneu 1996, leg. no. Distribution: Palaearctic, new re monidae 1998. Taxapad CD-Rom. cord for Scandinavia Received 5 October 2000, accepted 2 November 2000 DISCUSSION About 1100 different species of Ichneumonidae have been found in Norway so far (Yu 1999). This paper gives distributional records of 169 additional species. Although the number of known Norwe gian lchneumonidae is substantially increased by this publication, many more of these parasitoid wasps can be expected to occur in Norway. With respect to other Scandinavian countries such as Sweden with approximately 2,500 reported ich neumonid species (Yu 1999), we estimate that not more than 1/2 to 2/3 of the Norwegian ichneu monid fauna have been reported so far. Therefore, the Ichneumonidae form the most species-rich family of animals in Norway as well as in most other countries in the Palaearctic region (Gauld & Bolton 1988, Yu 1999). Most of the species mentioned in this paper are widespread in Europe and were previously found in other Northern countries. However, fifteen spe cies which were collected in the warmer South eastern and Southern parts ofNorway are reported from Scandinavia for the first time. Acknowledgements. For the collection and loan oftheir ichneumonid material we would like to thank all the collectors in Norway who generously donated their material for this study, especially Mr Thor Jan Olsen. References Gauld, I.D. & Bolton, B. 1988. The Hymenoptera. 332 pp. Oxford University Press, Oxford. Hinz, R. 1973. Beitriige zur Kenntnis der Arten der Ichneumoninae. Ent. Nachr. 17,97-105. 0kland, K.A. 1981. Inndeling av Norge til hruk ved 176 Norw. J. Entomol. 47, 177-181.2000 Aculeata of Norway. 3. Eleven species of bees new to Norway (Hymenoptera: Apoidea) 0istein Berg Berg, 0. 2000. Aculeata of Norway. 3. Eleven species of bees new to Norway (Hymenoptera: Apoidea). Norw. J. Entomol. 47,177-181. The following eleven species ofbees (Apoidea) are published from Norway for the first time: Colletes marginatus F. Smith, 1846, C. similis Schenck, 1853, Hylaeus annularis (Kirby, 1802), H. rinki (Gorski, 1852) (Colletidae), Andrenafulvida Schenck, 1853, A. ovatula (Kirby, 1802), Panllrginlls romani Aurivillius, 1914 (Andrenidae), Lasioglossum xanthopus (Kirby, 1802) (Halictidae), Dasypoda hirtipes (Fabricius, 1793) (Melittidae), Osmia aurllienta (Panzer, 1799) and Megachile lapponica Thomson, 1872 (Megachilidae). A total of 184 species of bees are now recorded from Norway. Key words: Hymenoptera, Apoidea, solitary bees, distribution in Norway. 0istein Berg, Kitl).' Kiellands vei 19 C, NO-1344 Haslum, Norway. INTRODUCTION erature. This is beyond the scope of the present work. Nearly all publications on Norwegian bees the last 60 years have been restricted to the family Some years ago a few Norwegian entomologists Apidae. The bumblebees (Apidae, Bombinae) took up an interest in the aculeate Hymenoptera. have been thoroughly worked by L0ken (1973, This article is number 3 in a planned series of 1984). The honeybee, Apis mel/ifera L., 1758 works on Norwegian aculeates. The previous two (Apidae, Apinae), has been intensively studied articles were published by Hansen (1995) and in several aspects. The non-apid bees, sometimes Hansen & Olmi (1996). called solitary bees, have been largely ignored. The only two complete checklists of Norwegian SPECIES LIST bees are old, being published by Siebke (1880) and Strand (1898). Regional lists have been pub Nomenclature follows Schwarz et al. (1996, p. lished by Soot-Ryen (1925) for North Norway 190). Biogeographic regions are given according and by Meidell (1934) for Rogaland. Additional to J. 0kland (1976) and EIS-squares according to records have been published by Sommerfelt K.A. 0kland (1981). The following abbreviations (1824-27), Zetterstedt (1838), Nylander (1852a, have been used: AF=Arne Fjellberg, LOH=Lars b), Sch0yen (1880, 1887), Sparre Schneider Ove Hansen, 0B=0istein Berg. (1895, 1909), Strand (1900, 1901, 1903, 1906, The author has identified specimens unless otherwise 1910), Friese (1902), Hellen (1933), Meidell noted in the text. The material is deposited in the pri (1945), L0ken (1958, 1967), Tkalcil(l974, 1975, vate collections of0B or AF unless otherwise noted. 1977a, 1977b, 1983), Greve & Hauge (1989) and Fjellberg (1993). A survey ofthis literature shows Colletidae that approximately 173 species ofbees have been published from Norway. To establish an exact Colletes marginatus F. Smith, 1846 number requires a revision of museum material AK Oslo: Gressholmen (EIS 28) 3 99, 2 eJ(} 10 to fully understand certain names used in old lit- 24 June 1992 on Vincetoxicum rossicum (Asclepia 177 Berg: Aculeata of Norway. 3. Bees new to Norway (Hymenoptera: Apoidea) daceae) leg. Emmy Gram, det. AF, coIl. Zoologi leg. 0B; TEY Nome: Ulefoss (EIS 18) 69931 cal Museum, Oslo; Brerum: Kalvoya (EIS 28) 3 July 1993 leg. 0B; TEl Bo: Verpe (EIS 17) I a aa 8 July 1995 leg. 0B; Asker: Nesoya (EIS 28) II June 1994 leg. 0B. I a 21 June 1992 leg. & coIl. LOH; VE Tjome: Moutmarka (EIS 19) 1 a 3 July 1992 leg. & det. Andrenidae AF; Tjome: Hellesmo (EIS 19) 2 99 26 July 1995 Andrena fulvida Schenck, 1853 and 1 a, 1 94 August 1996 leg. 0B. RES Eidskog: Leirsjoen (EIS 38) 2993 July 1991 C. similis Schenck, 1853 leg.0B. AK Brerum: Kalvoya (EIS 28) 1 96 July 1991 It is interesting to note that this species was also leg. & det. AF, 2 aa 8 July 1995 and 1 a 12 July recently published for the first time from Sweden 1999 leg. 0B; Brerum: Fleskumasen (EIS 28) 19 (Svensson, Erlandsson & Janzon 1990). 24 August 1998 on Achillea millefolium (Compo sitae) and 1 a 3 July 1999 leg. 0B; Asker: Hval A. ovatula (Kirby, 1802) (EIS 28) 1 9 2 aa from a row of cells found in (syn. albofasciata Thomson, 1870) ground 30 April 1995. An agricultural vehicle had VE Tjome: Hvasser, near Sonstegard (EIS 19) 1 cut a vertical profile in the soil and thus exposed 97 June 1993 leg. 0B. the cells. Pupae were kept indoors and hatched 30, 28 and 27 May 1995 respectively. Leg. 0B; 80 This species belongs to the subgenus Taeniandrena Royken: Kinnartangen (EIS 28) 1 9 II July 1993, (sensu Warncke 1968) that has very distinct white leg. 0B; Kongsberg: Labru (EIS 27) I 9 29 June bands across gaster. The other Norwegian species 1993 leg. Bjorn A. Sagvolden; VE Vale: Langoya of this subgenus are gelriae van der Vecht, 1927, (EIS 19) 1 9 Malaise-trap 8 July-2 August 1991 intermedia Thomson, 1870, lathyri Aitken, 1899 leg. LOH; Tjome: Moutmarka (EIS 19) I a 8 June and wilkella (Kirby, 1802). Niemela (1949) gives 1992 leg. & det. AF; Tjome: Sando (EIS 19) 1speci a key to these species. men 7 June 1992 leg. & det. AF; Tjome: Hvasser, Fyn (EIS 19) 2 aa 4 August 1996 on Tanacetum Panurginus romani Aurivillius, 1914 vulgare (Compositae) leg. 0B; Larvik: Molen (EIS OS Lillehammer: Smestad (EIS 54) I 9 14 June 11) I a 24 June 1992 leg. AF; TEY Porsgrunn: 1978 leg. Svein Svendsen, coIl. Tore R. Nielsen; Sandoya (EIS 11) 1916 July 1998 leg. 0B. STI Midtre Gauldal: Storen (EIS 87) 1 9 14 July 1993, leg. & coIl. Tore R. Nielsen; NTI Stjordal: Hylaeus annularis (Kirby, 1802) Vikan (EIS 92) 3 aa 21 June 1998, 2 99 1 a 15 VE Tjome: Moutmarka (EIS 19) 1 a 8 June 1992 July 1998, I 9 17 July 1998, leg. Lay & Rykken, leg. 0B; Tjome: Sando (EIS 19) 2 aa 7 June 1992 coIl. Scott Armbruster, Norwegian University of leg. 0B; Larvik: Oddane Sand (EIS 11) I 9 3 Au Science and Technology, Trondheim; NSI Grane: gust 1999 leg. 0B; AAY Arendal: Merdoy (EIS Trofors (EIS 115) 2 99 25 July 1946 leg. Tron 6) I a 13 June 1992 leg. 0B; Arendal: Flostaoya, Soot-Ryen, coil. Zoological Museum, Troms0. Arnevik (EIS 6) 3 99, I a 13 July 1997 leg. 0B; This is the first record of this genus in Norway, YAY Sogne: Lyngmyr (EIS 2) I 9 29 July 2000 and the only boreal non-apid bee species recorded leg.0B. in Norway. H. rinki (Gorski, 1852) Halictidae AK Brerum: Fleskumasen (EIS 28) I a 18 June 1999 leg. 0B; OS Gjovik: Redalen (EIS 54) I 99 Lasioglossum xanthopus (Kirby, 1802) July 1999 leg. 0B; 80 R0yken: Hyggen, Kinnar o Fredrikstad: Gansf0d/0ra (EIS 20) 1 930 Au tangen (EIS 28) I a 11 July 1993 leg. 0B; VE gust 1991, leg. Thor Jan Olsen. Lardal: Hukstmm (EIS 19) 39915 August 1991 178 Norw. J. Entomol. 47, 177-181.2000 Melittidae scribed so far, while he believes the total number of species could be as high as 30 000. Dasypoda hirtipes (Fabricius, 1793) () Rade: Aven (EIS 19) 5 eJcJ 9-11 August 1996, Both above mentioned species ofHylaeus belong leg. & colI. LOH; VE Tjome: Hellesmo (EIS 19) 4 to the subgenus Lambdopsis Popov, which is here ex; 26 July 1995,499 4 August 1996, leg. 0B; a reported from Norway for the first time. It is in colony with about 800 nests was established at Hel teresting to note that in the small material so far lesmo in August 1997 (AF pers. comm.); Tjome: examined, all records of H. annularis are from Hvasser, Nessletta (EIS 19) 194 August 1996, leg. localities very close to the sea. Most records ofH. 0B; also observed at Tjome: Sando (EIS 19) by rinki are from inland localities. AF. HINTS ON USEFUL LITERATURE Megachilidae A brief general introduction to solitary bees, in Osmia aurulenta (Panzer, 1799) Norwegian, is given by Fjellberg (1995). A very useful article by Nilsson & Svensson (1986) con VE Tjome: Sando (EIS 19) 1 male 16 June 199 L tains a lot of basic information needed ifone wis 299 30 May 19921eg. & colI. AF; Tjome: Hvasser, hes to start studying North European acu1eates. In near Sonstegard (EIS 19), abandoned shells of particular there is a long list of identification lit Cepaea nemoralis with the opening plugged by a erature. The most useful to start with is probably mastic of chewed leaves were collected on the 6 Landin (1971), which gives a key to the Swedish November 1991, leg. 0B. In April and May 1992 genera. Recently there have been published excel 5 aa of O. aurulenta emerged. Same locality: 19 lent keys in German by Scheuchl (1995, 1996) and 7 June 1993 leg. 08. Schmid-Egger & Scheuchl (1997). These contain many figures and should enable identification of Megachile lapponica Thomson, 1872 all Central European species ofthe families Andre AK Fet: Enebakkneset (EIS 29) 1 927 June 1990 nidae, Melittidae, Megachi1idae and Anthopho leg. 0B; BV Rollag: Rollag (EIS 35) 1 a 8 Sep ridae. All species published from Norway are in tember 1993 leg. Bjorn A. Sagvolden. cluded except Panurginus romani Aurivillius. The disadvantage of such comprehensive works em DISCUSSION ployed on the Norwegian fauna is that they con tain a large number of species not occurring in Including the species listed above, there are now Norway. Many excellent photographs and a lot of recorded some 184 species of bees in Norway. As information on biology can be found in Westrich will be seen from the species list, most ofthe ma (1990). The same can be said about two smaller terial examined for this study has been collected books by O'Toole & Raw (1991) and Muller et al. in the lowlands ofsoutheastern Norway. This area (1997). A standard reference work for bees will appears to have the highest species diversity of be Michener (2000) that contains, among other, non-apid bees in Norway. Most species of non keys to all genera and subgenera of the world. apid bees are known to be xerothermophilous, thus, the number of species increases towards the south. This is true both within Norway and fur Acknowledgements. For help with identification I wish to thank F. Gusenleitner, Austria (Andrena), Lars Noren, ther south. Austria is known to have a total of647 Sweden (Lasioglossum) and the late Gijs van der species ofbees (Schwarz et at. 1996), while Israel Zanden, the Netherlands (Megachile). For gift and loan has an estimated 1500-2000 species (O'Toole & ofmaterial I would also like to thank ScottArmbruster, Raw 1991). The highest diversity ofbees is found Emmy Gram, Lars Ove Hansen, Tore R. Nielsen, Arne in warm temperate areas, such as the Mediterra Nilssen (Zoological Museum, Tromso), Tor Jan Olsen nean area. Michener (2000) estimates that about and Bjorn A. Sagvolden. For useful comments and ad 17 000 different species of bees have been de- ditional records, thanks to Arne Fjellberg and Lars Ove Hansen. 179 Berg: Aculeata of Norway. 3. Bees new to Norway (Hymenoptera: Apoidea) References alium, comparatis speciebus Europ1e medi1e. Notis. Fjellberg, A. 1993. Rapport fra Pinsetreffen 1992 Saellsk. Faun. F!. fenn. Forh. 2,225-286. Tjome. Insekt-Nytt, 18 (1),11-12. 0kland, J. 1976. Utbredelsen av noen ferskvannsmus Fjellberg, A. 1995. Villbier - bestovning viktigere enn linger i Norge, og litt om European Invertebrate honning. Insekt-Nytt, 20 (1-2),41-43. Survey. Fauna (Oslo) 29, 29-40. Friese, H. 1902. Die arktischen Hymenopteren, mit 0kland, K.A. 1981. Inndeling av Norge til bruk ved Ausschluss der Tenthrediniden. Fauna Arctica 2, biogeografiske oppgaver - et revidert Strand-sys 439-498. tem. Fauna (Oslo) 34,167-178. Greve, L. & Hauge, E. 1989. Insekt- og edderkopp O'Toole, C. & Raw, A. 1991. Bees of the World. 192 faunaen pa myrer i Hordaland. I. Iglatjodn - Stord pp. Blandford, London. kommune. II. Sjoalemyr - Stord kommune. Rap Scheuchl, E. 1995. 11lustrierte Bestimmungstabellen der port terrestrisk okologi nr. 49. Zoologisk museum, Wildbienen Deutschlands und Osterreichs. Band I: Universitetet i Bergen. Schliissel der Gattungen und der Arten der Familie Hansen, L.O. 1995. Aculeata of Norway. I. Bethylidae Anthophoridae. 158 pp. Privately Published, Velden. (Hym., Apocrita). Fauna norv. Ser. B 42, 43-48. Scheuchl, E. 1996. Illustrierte Bestimmungstabellen der Hansen, L.O. & Olmi, M. 1996. Aculeata ofNorway. 2. Wildbienen Deutschlands und Osterreichs. Band II: Dryinidae and Embolemidae (Hym., Apocrita). Schliissel der Arten der Familien Megachilidae und Fauna norv. Ser. B 43,81-88. Melittidae. 116 pp. Privately Published, Velden. Hellen, W. 1933. Hymenoptera Aculeata aus Finmarken Schmid-Egger, C. & Scheuchl, E. 1997. Illustrierte in Norwegen. Notul. ent. 13,41-43. Bestimmungstabellen der Wildbienen Deutschlands Landin, B.-O. 1971. Insekter 2:2. Faltfauna. 672 pp. und Osterreichs unter Beriicksichtigung der Arten Natur och Kultur. Stockholm. der Schweiz. Band III: Schliissel der Arten der Loken, A. 1958. Pollination studies in apple orchards Familie Andrenidae. 180 pp. Privately Published, of Western Norway. Proceedings 10th Int. Congr. Velden. Entomol., 1956, vo!. 4, 961-965. Schoyen, WM. 1880. Bem1erkninger til H. Siebke's Loken, A. 1967. A stylopized female ofAndrenafucata Enumeratio insectorum Norvegicorum, Fasciculus Smith (Streps.-Hym.). Opusc. ent. 32, 93-96. V. Pars I (Hymenoptera phytophaga & aculeata), Loken, A. 1973. Studies on Scandinavian Bumble Bees ediditJ. Sp. Schneider. Forh. Vidensk.Selsk. Christi (Hymenoptera, Apidae). Norsk Ent. Tidsskr. 20, 1 ania, no. 10, 1-15. 218. Schoyen, WM. 1887. Supplement til H. Siebke's Enu Loken. A. 1984. Scandinavian species of the genus meratio insectorum Norvegicorum Fasc. V, Pars I Psithyrus Lepeletier (Hymenoptera: Apidae). Ent. (Hymenoptera phytophaga & aculeata). Forh. scand. Suppl. 23, 1-45. Vidensk.Selsk. Christiania for 1887, no. 5, 3-11. Meidell, O. 1934. Bier og humler i Rogaland. Stavanger Schwarz, M., Gusenleitner, F., Westrich, P. & Dathe, H.H. Museums Arshefte, 43. argang (1932-33), 85-131. 1996. Katalog der Bienen Osterreichs, Deutschlands Meidell, O. 1945. Notes on the Pollination of Melamund der Schweiz (Hymenoptera, Apidae). Entomo pyrum pratense and the «Honeystealing» of Hum fauna, Supp!. 8, 1-398. ble-bees and Bees. Bergens Mus. Arb. 1944. Nat. Siebke, H. 1880. Enumeratio Insectorum Norvegicorum. rekke no. 11, 1-12. Catalogum hymenopterorum continentem. Auctore Michener, C.D. 2000. The Bees of the World. 872 pp. defuncto H. Siebke, ed. 1. Sparre Schneider, 5 (1). Johns Hopkins University Press, Baltimore. A.W Broegger, Christiani1e. Miiller, A.. Krebs, A. & Amiet, F. 1997. Bienen. Mittel Sommerfelt, S.Chr. 1824-27. Physisk-oeconomisk europaische Gattungen. Lebensweise, Beobachtung. beskrivelse over Saltdalen i Nordlandene. Insecter. 384 pp. Naturbuch Verlag, Munich. Kong!. Norske Vid. Selsk. Skrifter, 2, 98-100. Niemela, P. 1949. Mitteilungen iiber die Apiden (Hym.) Soot-Ryen, T. 1925. Entomologische Notizen I. Hyme Finlands. 3. Die Untergattung Taeniandrena Hedi noptera aculeata und tubulifera aus dem nordlichen cke. Ann. Ent. Fenn. 15,101-120. Norwegen. Tromso Mus. Arshefter 1924,47, nr. 3, Nilsson, G.E. & Svensson, B.G. 1986. Handledning for 1-15. gaddstekelsamlare. Ent. Tidskr. 107, 151-166. Sparre Schneider, H.J. 1895. En entomologisk udflugt Nylander, W. 1852a. Supplementum Adnotationum in til Bardodalen og Altevand i juli 1893. Ent. Tidskr. Expositionem Apum Borealium. Notis. Saellsk. 16,225-248. Faun. F!. fenn. Forh. 2,93-107. Sparre Schneider, H.J. 1909. Hymenoptera aculeata im Nylander, W 1852b. Revisio synoptica Apum Bore arktischen Norwegen. Tromso Mus. Aarshefter 1906,29,81-161. 180 NolW. J. Entomol. 47, 177-181.2000 Strand, E. 1898. Enumeratio Hymenopterorum Norve gicorum. Ent. Tidskr. 19, 71-112. Strand, E. 1900. Et \idet bidrag til Norges entomologiske fauna. Ent. Tidskr. 20 (1899), 287-292. Strand, E. 1901. Orthoptera og Hymenoptera samlcde i 1900. Arch. Matern. Naturvidensk. 23 (6), 1-7. Strand, E. 1904. Hymenopterologisk bidrag til Norges fauna. Forh. Vidensk.Selsk. Christiania for 1903, no. 8, 1-8. Strand, E. 1906. Nye bidrag til Norgcs hymenopter- og dipterfauna. Nyt Mag. Naturvidensk. 44 (1906),95 104. Strand. E. 1910. Neue Beitriige zurArthropoden-Fauna Norwegens nebst gelegentlichen Bemerkungen tiber deutscheArten. I-V. Nyt Mag. for Naturvidensk. 48 (1910),307-354. Svensson, B.G., Er1andsson, S., & lanzon, L.-A. 1990. Catalogus Insectorum Sueciae. Hymenoptera, Apoi dea. 2. Andrenidae and Halictidae. Ent. Tidskr. 111, 47-52. Tka1cU, B. 1974. Revision und K1assifikation der bisher zur Untergattung Hoplosmia Thomson gestellten Anthocopa-arten (Hymenoptera, Apoidea, Megachi lidae). Acta Entomol. Bohemoslov. 71, 114-135. Tkalcu. B. 1975. Revision der europiiischen Osmia (Chalcosmia)-Arten der fiilviventris Gruppe (Hy menoptera: Apoidea: Megachi\idae). Vest. csl. Spol. zool. 39,297-317. Tkalcu. B. I977a. Die Osmia-Arten der Untergattung Neosmia Tkalcu (Hymenoptera, Apoidea, Megachi lidae). Acta Entomol. Bohemoslov. 74, 85-102. Tkalcu. B. 1977b. Taxonomisches zu einigen paliiarkti schen Bienenarten (Hymenoptera: Apoidea). Vest. csl. Spol. zool. 41, 223-239. Tkalcu. B. 1983. Die europiiischen Osmia-Arten der Untergattung Melanosmia (Hymenoptera, Apoidea, Megachi1idae). Vest. csl. Spol. zool. 47,140-159. Warncke, K. 1968. Die Untergattungen der westpalii arktischen Bienengattung Andrena F. Mems Estud. Mus. zool. Univ. Coimbra 307,1-110. Westrich, P. 1990. Die Wildbienen Baden-Wtirttem bergs. 2 vols., 972 pp. 2nd ed. Ulmer, Stuttgart. Zetterstedt, l.W. 1838. Insecta Lapponica descripta. 1139 pp. Voss, Leipzig. Received 7 March 2000, accepted 25 October 2000 181 Short communication The genus Pteromicra Lioy, 1864 the provinces ON and NSI by Greve & Rozkosny (Diptera, Sciomyzidae) in Norway (1981). No additional records ofthis species were noted by Rozkosny (1984). Lita Greve Collection ofSciomyzidae for several decades has turned up very few new records ofthe genus Pte romicra, but no new species for Norway. All this Greve, L. 2000. The genus Pteromicra Lioy, 1864 (Di new material is deposited in Zoological Museum, ptera, Sciomyzidae) in Norway. Norw. J. Entomol. 47, University ofBergen. Regional abbreviations are 182-183. given in accordance with 0kland (1981). The distribution ofthe genus Pteromiera in Norway is NEW RECORDS reviewed. Two species Pteromicra angustipennis (Stae ger, 1845) and Pteromiera glabricula (Fallen, 1820) Pteromicra angustipennis (Staeger, 1845) have been recorded from southern Norway north to RY Finnoy: Kyrkoy (EIS 14) 22 June 1986 19 Nordland province and both are rare. leg. & det. T. Jonassen (ZMB). HOI Kvam: Grav Key words: Pteromicra, Sciomyzidae, Diptera. dal, Svevatn (EIS 31) Malaise trap 29 July - II Sept. 1997 19 leg. J. Skartveit. SFY Hyllestad: Lita Greve, University a/Bergen, Zoological Museum, Botnen (EIS 48) Malaise trap 21 May - 21 June Museplass 3, N-5007 Bergen-Univ, Norway. 1999 IeJ 1921 June - 25 July 1999 3eJeJ 399 leg. Lita Greve & Liv Sognes. P angustipennis has previously been recorded INTRODUCTION from ON & NSI, and is here reporteed new to RY, RozkoSnY (1984) listed five species of the genus HOI and SFY. Pteromicra Lioy, 1864 in his review of the Scio myzidae of Fennoscandia and Denmark. Of these Pteromicra glabricula (Fallen, 1820) Pteromicra oldenbergi (Hendel, 1902) is a rare HES Yang: Bla:stad, Hedmark Distrikthoyskole species which has only been recorded from the most 17 June 1996 19 leg. L. Greve. YAY Flekkefjord: south-easternly parts ofFinland and Russian Kare Hidra, Dragoy 8-15.Aug.1992 IeJ leg. A.- J.Nilsen len and should not be expected to occur in Nor (ZMB). way. Two other species have a southern distribution P glabricula has been recorded from ON, VE & in Fennoscandia, P leucopeza (Meigen, 1838) is NSI, and is here reported new to HES and VAY. recorded from south-eastern Sweden, southern Fin land and Denmark, P pectorosa (Hendel, 1902) is recorded only from south-eastern Sweden and in DISCUSSION Denmark, and not from Finland (RozkoSnY, 1984). The genus Pteromicra in the tribe Sciomyzini is a One species, P glabricula (Fallen, 1820), was re well defined genus which contains generally small corded from Norway by Zetterstedt (1838). The species with dark brown to black and yellow distribution ofP glabricula was given as the prov ground colour. The anterior orbital seta is often inces ON, VE and NSI by Knutson & Berg (1971 ), short and the cheeks very narrow. The propleuron and they listed only one record from each prov carries a rather weak, but distinct seta above base ince. According to RozkoSnY (1984) P glabricula of fore coxa, the anal vein reaches the posterior is recorded «in Norway several records from ON margin of the wing. Wings are either hyaline or to NSI», but the correct number ofrecords are the completely infuscated. The fore tibia has one pre three mentioned by Knutson & Berg (1971). apical seta. They are not very difficult to identify and should therefore not have been overlooked or A second species, Pteromicra angustipennis (Stae mistaken for other species or genera of Sciomy ger, 1845), was recorded as new to Norway from zidae. 182 Norw. J. Entomol. 47, 182-183.2000 Based on the records presented here both P angus Book review • Bokanmelde/se tipennis and P glabricula must be considered rare species in Norway. The records of both species are few and widely scattered. Aarvik, L., Berggren, K. &Hansen, L.O. (red.) 2000. Catalogus Lepidopterorum Norvegiae. Acknowledgements. I am grateful to Terje Jonassen 192 pp. Lepidopterologisk arbeidsgruppe, Zoo who collected and determined the specimen of Ptero logisk Museum, Universitetet i Oslo og Norsk micra angustipennis from Rogaland, and to the follow institutt for skogforskning, As, Norway. ISBN 82 ing persons who have provided material: John Skartveit 995095-1-3. [NOK 200,- (paperback) NOK 250, and the research programme «Forest ecology and mul (hardback), bestilles fra Insektavdelingen, Zoo tiple use»,Alf-Jacob Nilsen, Hidra and Liv Sognes, Hyl lestad. logisk Museum, Sarsgate 1, 0562 Oslo]. Sa har den endelig kommet, katalogen over Nor References ges sommerfugler, nesten 70 ar etter at den forrige Greve, L. & RozkosnY, R. 1981. Three species of Scio fullstendige oversikt over Norges sommerfuglfau myzidae (Dipt.) new to Norway, with a note on Ecti na sa dagens Iys. Den gang var 1567 arter pavist nocera borealis Zetterstedt 1838. Fauna norv. Ser.B fra landet. Med Lepidopterologisk arbeidsgruppe 28, 100 - 10 1. (LepArb) sitt kjempeverk, kan det na dokumen Knutson, L.v. & Berg, C.O. 1971. The Malacophagous teres at heIe 2123 arter av sommerfugler er pavist Flies ofNorway (Diptera, Sciomyzidae). Norsk ent. i Norge. Katalogen er basert pa det aller meste av Tidsskr. 18, 119 - 134. tilgjengelig sommerfuglmateriale i offentlige og 0kland, K.A. 1981. Inndeling av Norge til bruk ved biogeografiske oppgaver - et revidert Strand-sys private samlinger i Norge, til sammen ca. 100 000 tem. Fauna (Oslo) 32, 167 - 178. registreringer. Mange feilkilder er eliminert ved RozkoSnY, R. 1984. The Sciomyzidae (Diptera) of Fen at hvert enkelt individ er kontrollbestemt av Lep noscandia and Denmark. Fauna ent. Scan. 14, 1 Arbs medlemmer. Deler av dataene er ogsa til 224. gjengelig via intemett, hvor landsdekkende utbre delseskart for aile arter samt oversikter over hvilke Received 5 June 2000, arter som er funnet i landets kommuner er tilgjen accepted 15 August 2000 gelige. Katalogen har bade norsk og engelsk tekst og har et lekkert design som gjor boken bade innbydende og tilgjengelig for en bred gruppe av lesere. Inn ledningen gir en oppsummering av sommerfugl forskningen i Norge fra midten av 1700-tallet og fram til i dag med LepArbs registreringsarbeid. Systematikken og nomenklaturen som er brukt i katalogen er svcert godt oppdatert og blir grundig gjennomgatt. De mange bidragsyteme takkes til slutt i innledningen. I sum er innledningsteksten informativ og nyttig, men kunne vcert noe mer gjennomarbeidet. Det forekommer bl. a. bastante pastander som opplagt kan diskuteres, og i tillegg er tituleringen av bidragsyteme noe inkonsekvent. Selve katalogen er bokens hoveddel og bestar av en sjekkliste for aile Nordens sommerfugler. Den norske utbredelsen til aile arter er angitt etter det reviderte Strand-systemet som deler landet inn i 37 regioner basert pa fylkes- og kommunegrenser. 183 Book review· bokanmeldelse Lesbarheten er sv 184 Norw. J. Entomol. 47, 185-195.2000 Distribution of Trichoptera in 0vre Heimdalen, Jotunheimen Mountains, Norway Sigve Reiso & John E. Brittain Reiso, S. & Brittain, lE. 2000. Distribution ofTrichoptera in 0vre Heimdalen, Jotunheimen Moun tains, Norway. Norw. J. Entomo!. 47. 185-195. The distribution oflotic trichopterans was studied in different vegetational zones, ranging from 1090 m a.s.! to 1550 m a.s.!, in the 0vre Heimdalen area in the Jotunheimen Mountains of southern Nor way. Five species were recorded in the mid-alpine zone, eleven species in the low alpine zone, and twenty-four species in the subalpine zone. In the mid-alpine zone the community was dominated by Apatania species and Apatania zonella was the only species found at 1550 m a.s.!. The dominant feeding modes were scrapers in the mid-alpine zone, and both shredders in the low alpine and subalpine zones. Temperature and food were important factors influencing species distribution. Many common running water species such as Potamophylax cingulatus. Halesus digitatus and Rhyacophila nubila had their altitudinal distribution limit in the low alpine zone. Filter feeding polycentropodids were found in lake outlets in the subalpine and low alpine zones. In the outlet of the small lake Bliltjern, in the mid-alpine zone, no filter feeding species were found. Many species we recorded in the subalpine zone had low abundance, indicating that these species lived at their altitudinal limit in this area. The first axis of DCA (Detrended Correspondence Analysis) clearly showed a gradient ofspecies in the 0vre Heimdalen area. The analyses indicated the species changes to be most distinct at the transition from the low alpine to the mid-alpine. Keywords: Trichoptera, distribution, vegetation, alpine zones. Sigve Reiso, Department ofBiology and Nature Conservation, Agricultural University ofNorway. N-1432 As, Norway. [Present adress: Ragna Nielsensvei 12. N-0592 Oslo, Norway]. John E. Brittain, Zoological Museum, University ofOslo. Sars gate 1, N-0562 Oslo, Norway. INTRODUCTION Heimdalen (Lillehammer & Brittain 1978) have been investigated. The Trichoptera are a large and diverse order of aquatic insects numbering about 10,500 described Several Norwegian studies stress the importance species world-wide (Solem & Gullefors 1995). of temperature as a major factor influencing life North-west Europe has about 250 species (Ander cycles and distribution of freshwater inverte sen & Wiberg-Larsen 1987). Due to their ability brates (Andersen & Tysse 1984, Brittain 1978, to produce silk, they can exploit a variety ofhabi 1983,1990, Eikeland 1982, Lillehammer 1974, tats and feeding modes. Lillehammer et a1. 1989, Sand 1996 and Solem 1985b). Solem (1985b) and Lillehammer & Previous studies on distribution of Norwegian Brittain (1978) compared the distribution of species have mostly been carried out in lowland aquatic insects in different vegetational zones. areas (Solem 1985b). Studies ofthe vertical dis From these studies and other studies on tribution of Trichoptera in mountain areas are Plecoptera (Lillehammer 1974, 1975, 1984), it sparse, although Dovrefjell (Solem 1985b), is clear that riparian vegetation is an important Hardangervidda (Andersen 1979) and 0vre 185 Reiso & Brittain: Distribution of Trichoptera in Jotunheimen Mountains, Norway factor influencing aquatic insect distribution in altitude as a result ofreduction in temperature and addition to temperature. allochthonous inputs. The aim of the study was to describe and under take a quantitative analysis of the distribution of Site description trichopteran species at altitudes ranging from 1095 The area investigated was the catchment area of m a.s.! in the subalpine zone to 1550 m a.s.! in the the subalpine lake, 0vre Heimdalsvatn (61 °25'N, mid-alpine zone in the 0vre Heimdalen area. It was g052'E), together with an adjacent stream catch hypothesised that species richness decreases with ment, Flybekken, on the eastern slopes of the Figure 1. Map of the catchment of 0vre Heimdalsvatn and the adjacent stream catchment, Flybekken. Altitudes in m a.s.1. 186 Narw. J. Enlamal. 47, 185-195.2000 Jotunheimen Mountains in southern Norway (Fig Heimdal area has long winters and the lake 0vre ure 1). The bedrock in the area consists almost Heimdalsvatn is ice-covered from mid-October to entirely ofbasic Precambrian rocks, which are part early June. The annual mean air temperature close of a large Caledonian thrust complex known as to the lake is estimated to be -1.2 DC, with a yearly the «Jotundekke» (Skjeseth & Kloster 1978). precipitation of 800 mm (Johannessen 1978). The area has a subarctic or tundra microclimate This study was mainly carried out in five streams, (Johannesen 1978). The northern side ofthe 0vre Brurskardbekken, Sandbakkbekken, Lektorbek Heimdalen valley has a favourable combination ken, Sorbekken and Flybekken (Figure 1). Sev of slope angle and radiation exposure allowing eral other small streams in the catchment area were subalpine birch forest to reach an altitude ofabout also sampled but less intensively. A wide range of 1200 m a.s.!. The southern side of the valley is sites were sampled to represent different vegeta dominated by willow scrub, with only a few scat tion belts between the elevations 1090 and 1550 tered birch trees. The vegetation along the stream m a.s.!. Three lake outlets were sampled, the out Flybekken is mostly willow and heath vegetation let of 0vre Heimdalsvatn in the subalpine zone up to 1200-1300 m a.s.!. At altitudes above 1300 (1090 m a.s.1), Brurskardtjern at the upper limit m a.s.! grasses dominates the vegetation. ofthe low alpine zone (1306 m a.s.1) and BIMjem in the mid-alpine zone (1465 m a.s.1). Following the vegetation zone definitions of 0sthagen & Egelie (1978), the low alpine vegeta Water temperatures were recorded by digital log tion occurs between 1200 and 1300-1400 m a.s.!. gers in the main inlet and outlet of 0vre Heim and the mid-alpine up to about 1800 m a.s.!. (Fig dalsvatn. The temperature data showed a rapid ure 1). The dominating vegetation types in the increase in temperature after the ice disappeared subalpine zone are mesotrophic birch forest and in early June, reaching over 10°C in the inlet and willow thickets, bilberry heath in the low alpine over 12 °C in the outlet in July/August. Water tem zone and heath, grass vegetation and snowbed peratures began to fall in late August approaching communities in the mid-alpine zone. o°C during October (Figure 2). The supply ofallochthonous matter to freshwaters is significant in the subalpine zone, decreasing in METHODS AND MATERIAL the low alpine zone and mid-alpine zone The trichopteran material was collected on seven (Lillehammer 1974, Larsson & Tangen 1975). The occasions during the summer of 1998. Two sam ------Outlet 14, - •••• -- --- _. Inlet 12 (J 10 Gl.. :,,::\,,:J\., ", ~ o I.,.,'" ./\.' ,", .. \ " " ' '. .' t 8 '~ :~ ;\"',;~ III ... .: ";,:':" :'\ ;-, -.. 6:' "', ::: .. '." ': ';..... \ Gl .\ I \,: ~,~. Q. 6 E '":; : \: , ':k\ Gl : o i I 21/5 416 18/6 2/7 16/7 30/7 13'8 27/8 10'9 24/9 Figure 2. Mean daily water temperatures (DC) in the outlet and inlet of 0vre Heimdalsvatn during 1998. The inlet temperature was recorded in Brurskardbekken, 250 m upstream of the lake. 187 Reiso & Brittain: Distribution of Trichoptera in Jotunheimen Mountains, Norway pIes were taken both during June and July then Adults were identified using Malicky (1983). No once in August, September and October. attempt was made to distinguish larvae of the closely related species Potamophylax cingulatus Larvae and pupae were collected by a kicking tech and P latipennis or Halesus digitatus and H. radia nique (F'rost et al. 1971) (30x30 em net, mesh size tus. Due to similarities in morphology they are 350 /-lm) and hand picked from the stone substra refered to as Potamophylax spp. and Halesus spp., tes. Adults were collected in three malaise traps, respectively. All the species were grouped into three one ultraviolet light trap and by sweep netting in categories based on their feeding habits (scrapers, the vegetation along streams. Malaise traps were shredders or predators/filter feeders), according to all placed in the subalpine zone, two close to the Solem & Gullefors (1995), Solem (1969, 1985a, lake 0vre Heimdalsvatn, and one in the birch for pers. com.). est by the stream Brurskardbekken at 1130 m a.s.l. An ultraviolet light trap was placed close to the Detrended Correspondence Analysis (DCA) was outlet of0vre Heimdalen. Sweepnet samples were used to illustrate the altitudinal gradient in spe taken at all altitudes when sampling the streams. cies composition using the ordination analysis pro gramme, CANOCO (Ter Braak 1995). A «down All samples were preserved in 70% alcohol in the weighting of rare species» option of CANOCO field, sorted and identified in the laboratory. The was used so that species that occured at few sites larvae were identified to genus using Solem & were given low weighting to minimise their influ Gullefors (1995) and to species using Edington & ence on the analysis. The aim of ordination is to Hildrew (1981), Hiley (1976) and Solem (l985a). arrange points such that those that are close to- Table 1. Species in different vegetation zones grouped according to their feeding habits. Zone Scrapers Shredders Predators Mid-alpine A. zonella C. vil/osa A. muliebris L. coensous A. hispida Low alpine A. hispida Potamopylax spp. Potamopylax spp. A. stigmatella Halesus spp. R. nubila E. dalecarlica P conspersa C. vil/osa P flavomaculatus L. coensous Subalpine A. stigmatella Potamopylax spp. Potamopylax spp. A. hispida A. lapponicus R. nubila C. vil/osa P conspersa Halesus spp. P flavomaculatus L. algosus H. clathrata L. bipunctatus A. obsoleta L. borealis L. coenosus L. extrincatus L. femoralis L. f1avicornis L. nigriceps L. subcentralis M. lateralis M. sequax H. c/athrata A. obsoleta 188 Norw. J. Entornol. 47, 185-195.2000 gether correspond to sites that are similar in spe RESULTS cies composition and those that are far apart cor Subalpine zone respond to sites that are dissimilar in species com position. The data was analysed as species pres Twenty-four species were recorded in the subal ence/absence in ten different 50 m altitudinal zones, pine zone (Figure 3). Shredders were clearly the from 1050 m a.s.! to 1550 m a.s.! (Figure 3). most dominant feeding group (Table 1). The nine Limnephilidae spp.; Asynarchus lapponicus, Mic ropterna lateralis, Micropterna sequax and Oxy ethira jrici were only found in low numbers as adults close to the lake, 0vre Heimdalsvatn. Pfla vomaculatus (36.5 %) dominated the malaise trap catches in the outlet area of 0vre Heimdalsvatn. Vegetation zones: I sub alp. II low alp. I! mid-alp. I Altitudinal zones: 1 2 3 4 5 6 7 8 9 10 Hydroptilldae Oxyethira frici Klepslek, 1891 Ryacophllidae - Rhyecophile nubile (Zetterstedt, 1840) Polycetropodldae Pfectrocnernie consperse (Curtis, 1834) ~ ~ Polycentropus f/evorneculstus (Pictet, 1834) ~ - --~ Phrygaeldae Hegenelle clethrsfa (Ko/eneti. 1848) ... Agrypnia obsolete (Hagen, 1864) Limephilidae - Apetanie hispide (Forsslund, 1930) ~ Apstenie mullebris McLechlan, 1866 - - Apatania stigmatelle (Zetterstedt, 1840) 1000. - ~ Apstanie zonelle (Zetterstedt, 1840) - Asynerchus lepponicus (Zetterstffdt, 1840) I- Cheetopteryx villose (Febricius, 1798) ... Ecclisopteryx delf1Cerlica Ko/enati, 1848 -- -- H./esus digitetus (Schrenk, 1781) ...~ - Ha/esus rediatus (Curtis, 1834) I - Umnephilus e/gosus (McLachlan, 1868) Limnephilus bipunctatus Curtis, 1834 - Umnephilus borealis (Zetterstedt, 1840) - -I Limnephilus coenosus Curtis, 1834 Umnephilus extrincetus McLachlen, 1865 ... - Umnephilus femore/is Kirby, 1837 I- Umnephilus r1avicomis (Febricius. 1787) ~ Umnephilus nigriceps (Zelterstedt, 1840) I- Umnephilus subcentra/is Brauer, 1857 I Microplema/atere/is (Stephens, 1837) I- Micropteme sequlix McLachlan, 1875 I- Potemophylax cingulatus (Stephens, 1837) Potamophylax letipennis (Curtis, 1834) I Elevation m a.s.l 1050 1150 1250 1350 1450 1550 Figure 3. Distribution of Trichoptera in different vegetational and altitudinal zones in the f2Jvre Heimdalen area. 189 Reiso & Brittain: Distribution of Trichoptera in Jotunheimen Mountains, Norway Rhyacophila nubila and P cingulatus larvae and stream, Flybekken, in this zone. Limnephilus coen adults were collected in most of the localities in sous adults were also colIected in a sweep net sam this zone and were the dominant species in ma ple close to the outlet of Bmrskardtjem. laise traps in the inlet area of0vre Heimdalsvatn. Halesus digitatus adults were also common in this Mid-alpine zone area. Adults of C. villosa were recorded in ma laise traps close to Bmrskardbekken. A few P Five species were collected in the mid-alpine zone conspersa larvae were found in the lower part of (Figure 3). The dominant feeding group was scrap the stream, Bmrskardbekken. ers (Table 1). Apatania zonella was collected both in Lektorbekken and Sandbakkbekken. It was the only species found at 1550 m a.s.1. However, the Low alpine zone species was not recorded in the Flybekken area Ten species were recorded in the low alpine zone where A. hispida was the dominant Apatania spe (Figure 3). The dominating feeding groups were cies. A. hispida was not found at the same alti shredders and predators/filter feeders (Table I). tudes in either Lektorbekken or Sandbakkbekken. Rhyacophila nubila and Potamophylax spp. lar Chaetopteryro; villosa larvae were quite common in vae were both found at most sampling sites in this smalI streams and springs in this zone, occuring in zone, being common in streams up to about 1300 the inlet streams of Brurskardtjern and in several m a.s.\. Halesus spp. larvae were not found at alti springs feeding Lektorbekken and Flybekken. It tudes higher than 1095 m a.s.1 on the south side of was found up to 1465 m a.s.! in the outlet of Bla the lake 0vre Heimdalsvatn. However, in the stre tjem. Apatania muliebris larvae were found in one am Bmrskardbekken (\285 m a.s.\.) larvae were spring in the headwaters ofLektorbekken (1460 m found in an area with large amounts of al1o a.s.1). Limnephilus coenosus was found in springs chthonous material. in the mid-alpine zone up to 1460 m a.s.\. Chaetopteryx villosa larvae were abundant in the inlet area of Sorbekken, close to 0vre Heimdals DCA analysis vatn. (1095 m a.s.1). Limnephilus coensous lar vae were found in a marshy area in the outlet ofa The results ofthe DCA analysis are summarised in small lake at the headwaters of Sorbekken. One Figure 4. The first and second ordination axes ex single E. dalecarlia larva was found in Flybekken plained 60.5 % and 21.1 %, respectively, ofthe to at 1200 m a.s.1 in an area with scattered willow tal variability (Table 2). Species like A. zonella, C bushes. villosa and L. coenosus that were found at high alti tudes in the mid-alpine zone showed high coordi Polycentropus .flavomaculatus and Plectrocnemia nate scores on the first axis. In contrast, species like conspersa larvae were found in the outlet of the L. nigriceps and L. algosus, only found in the lake, Bmrskardtjem. Plectrocnemia conspersa lar subalpine zone, showed low coordinate scores. The vae and pupae were also found in high densities in second axis was more difficult to interpret. Species a slow flowing reach ofBrurskardbekken at about like P .flavomaculatus, c. villosa and P conspersa 1285 m a.s.\.A. hispida. A. stigmatella andL. coeno showed high coordinate scores, whileA. hispida and sus larvae were found on several occasions in the A. stigmatella showed low coordinate scores on the Table 2. Summary of the DCA analysis. Axes 2 3 4 Total inertia Eigen values: 0.605 0.211 0.070 0.005 1.1968 Lengths of gradient: 4.314 2.301 2.411 2.588 190 Norw. J. EntomoJ. 47, 185--195. 2000 second axis. The altitudinal zone points show a pat DISCUSSION tern ofincreasing altitude along the first axis. How Studies of the distribution of Trichoptera in the ever, zone 6 does not fit into this pattern, showing 0vre Heimdalen area showed differences in num lower coordinate scores than expected (Figure 4). ber of species captured in the various alpine veg Zones 1-4 (1350-1550 m a.s.!) and zones 5-10 etation zones. Studying stoneflies in the same area, (1050- 1350 m a.s.!) tended to form two separate Lillehammer (1984) found temperature and food groups along the first axis, indicating a change in to be the most important factors influencing the species composition around 1350 m a.s.l. reduction of species with altitude. Species like Apatania zanella, A. muliebris and Chaetapteryx villasa, found in the mid-alpine zone, can adapt to different temperature regimes by switching from 3.5 eA obs. eP/la evil P. con. e e A. zan. 10 8 • 6 + + Hales. spp. e 2 5 + + 7 ,; L cae. • + 9 e* Potam. spp. 3. R. nub. .A. sli. 4 e e A. mul A. his. e E dol -2.5 -1.0 4.5 * =B. bip., L. bar., L. nig., L. sub., H. cia., O. fri., L. fla., A. lap., L. lat., L. ext., L. fern., L. alg., M. seq. & M. lat. Figure 4. DCA ordination diagram of trichopteran species recorded in ten altitudinal zones, each occu pying 50 m from zone 1 (1050-1100m a.s.l) to zone10 (1500-1550m a.s.I). Altitudinal zones are marked with crosses, while species are marked with circles. The scale marks are multiples of the standard deviation. 191 Reiso & Brittain: Distribution of Trichoptera in Jotunheimen Mountains, Norway univoltine to semivoltine life cycles (Solem the Arctic archipelago ofSvalbard (Solem 1985b), 1985b, Gislason 1981 and Andersen & Tysse indicating this species to be well adapted to ex 1984). However, other species such as Potamo treme conditions. The distributions of A. zanella phylax spp. and probably Halesus spp. found in and A. hispida did not overlap in the 0vre Heim the low alpine and subalpine zones show constant dalen area. The Flybekken area was dominated life cycle length, being univoltine throughout their by A. hispida, while the streams to the north of distributional range (Higler & Solem 1986, Solem the lake, 0vre Heimdalsvatn, were dominated by & Gullefors 1995). The effect of food is related A. zonella. Aagaard et al. (1989) found A. hispida to reduction in both autochthonous and especially to be the dominating Apatania species in the allochthonous production at high altitudes (Lille subalpine zone, while A. zonella dominated the hammer 1984, Petersen et al. 1995). Lillehammer alpine zone. Some environmental factor must fa & Brittain (1978) found change in species distri vour A. zonella on the north side ofthe lake. How bution in 0vre Heimdalen to be paralleled by ever, the south facing slopes of thc north side of changes in terrestrial vegetation for Plecoptera, the lake have a higher radiation exposure, and Ephemeroptera and Trichoptera. They found re conditions should be more favourable than the duction in species to be greatest among the detri Flybekken area on the south side. tus feeders when comparing the subalpine and In contrast to Apatania spp., Chaetapteryx villosa mid-alpine zones. However, production of plant show other characteristics (Andersen & Tysse material is closely related to temperature, mak 1984), it reproduces sexually and it is a shredder. ing it difficult to separate these two factors. However, like Apatania spp. it is able to adapt to In the mid-alpine zone only limnephilids occurred low temperatures and short ice-free period by (Figure 3). Solem (l985b) also recorded this pat switching to a semivoltine life cycle at high alti tern from his study in the Dovrefjell Mountains tudes (Andersen & Tysse 1984). C. villosa adults in central Norway. Three ofthe five species found arc also well adapted to supercooling, and are bet in the mid-alpine zone ofthe 0vre Heimdalen area ter able to survive low temperatures when emerg belong to the genus Apatania. The three Apatania ing than other species found at lower altitudes likc species are parthenogenetic, have preference for R. nubila and L. borealis (Solem 1985b). small streams and are scrapers (Solem 1985a). In the low alpine zone shredders became a more Having a semivoltine life cycle strategy at high dominant feeding group (Table I), demonstrating altitudes (Solem 1985a), these species are well the importance of allochthonous riparian inputs. adapted to the harsh environment and short sum According to Solem (l985b), R. nubila, P mers in the mid-alpine zone. In species having both cingulatus, P conspersa and E. dalecarlica are sexual and parthenogenetic reproduction, parthe common species in the low alpine and subalpine nogenesis dominates at high altitudes and latitudes zones. This agrees with the observations in 0vre (Butler 1984) where swarming is difficult. Heimda1en, although only one individual of E. Feeding on periphyton, the Apatania species can dalecarlica was recorded. Therfore, the low al survive in stream reaches where the terrestrial pine zone seems to be the upper altitudinal limit vegetation is sparse and allochthonous inputs are for many common lotic species. limited. In such areas, light is not limited, at least Filter feeders are common in lake outlets (Allan during summer. This enables benthic diatoms and 1995), but in the outlet area of Blatjcrn (1465 m algae to dominate the entire stream. In an alpine a.s.!) in the mid-alpine zone no filter feeding stream in northwestern Iceland, Odinsson (1988) trichopterans were found. However, both the fil found the macroinvertebrate benthic community ter feeding P flavomaculatus and P conspersa to be dominated by scrapers. occurred in the outlet of Brurskardtjem (1306 m Apatania zonella was the only species found above a.s.!) at the upper limit of the low alpine zone. 1500 m a.s.! (Figure 3). The species is also the Andersen (1979) found neither Pflavomaculatus only trichopteran recorded as far north as 80° on nor P conspersa above 1275 m a.s.! in his study 192 NoTW. J. Entomol. 47, 185-195.2000 oftrichopterans from the Hardangervidda moun tella as a species living below the tree line and tain plateau. Solem (1985b) only recorded the two being univoltine throughout its range. However, species in the subalpine zone in the Dovrefjell in Flybekken the species was recorded above the Mountains. Filter feeding trichopteran species are tree line in the low alpine zone. therefore probably at or near their altitudinal limit The first axis in the DCA analyses shows clearly at 1306 m a.s.l in the outlet of Brurskardtjern. a gradient of species in the 0vre Heimdalen area Lillehammer (1984) found potential prey species (Figure 4). The first axis describes environmental like chironomids and Baetis spp. to be numerous factors that changes with altitude. The analyses at a 1400 m a.s.! in a small lake on the nearby indicated the species change in the 0vre Heim Valdresflya. This lake resembles the lake, Bliitjern. dalen area to be most obvious in the transition Temperature and not food seems therefore to be from low alpine to mid-alpine vegetation. The se the limiting factor for filter feeders. cond axis, however, is more difficult to interpret In the subalpine zone, the number of species is when lacking environmcntal or physical data in doubled compared to the low alpine zone (Figure the analysis. Apatania muliebris, A. obsoleta, E. 3). However, many species had low abundance, darlecarlia and P flavomaculatus are at the edge indicating that many species live at their altitudinal ofthe diagram. Such species have little influence limit. In the subalpine zone the deciduous ripari on the analysis, lying there because they have few an vegetation influence the streams both by shad records (Ter Braak 1995). However, species at the ing and organic inputs. This is reflected in the do cdgc ofthe diagram can also be rare because they minance of shredders. Larsson & Tangen (1975) prefer particular environmental conditions or habi have shown that the inlet streams of 0vre Heim tats. This seems to be the case for P flavomacu dalsvatn transport large amounts ofallochthonous latus, restricted to lake outlets. material, explaining why shredders like P cingu The length of the axis is expressed in multiples of latus, H. digitatus and C. villosa were common in the standard deviation. Sites that differ four stand these areas. ard deviations in scores can be expected to have Limnephilus borealis and Asynarchus lapponicus no species in common (Ter Braak 1995). The length occur in ponds or lakes (Andersen 1979). The two of the first axis is estimated to 4.314 s.d., indicat species were only recorded as adults and might ing that zones I (1050-11 00 m a.s.!) and 10(1500 therefore originate from the lake, 0vre Heimdals 1550 m a.s.!) have no species in common. Apatania vatn, and not be part of the stream benthos. The zonella, the only species found in zone I, had a filter feeding P flavomaculatus was found in high restricted mid-alpine distribution in 0vre Heim densities, being the dominant trichopteran species dalen. However, the species has previously been in the outlet of0vre Heimdalsvatn (Lillehammer recorded down to sea level (Andersen 1979) and and Brittain 1978, Eikeland 1982 and Reiso 1999). might be found at altitudes similar to zone 1. How This indicates the importance of drift out of the ever, most species found in zone 1are likely to reach lake for the fauna in outlet areas (Larsson et al. their altihldinallimit before reaching zone 10. 1978). Adult Halesus radiatus was recorded close According to the first axis of the analysis, zone 5 to the outlet of0vre Heimdalsvatn, indicating that (1250- 1300 m a.s.!) tends to be closer to low alti the Halesus larvae found in the inlet streams were tude conditions This could be due to the presence H. digitatus larvae. In the Dovrefjell Mountains, of Halesus larvae at this site. Larvae were found Solem (1985b) found a similar pattern. He found in Brurskardbekken at this altitude in a pool rich H. radiatus to be common in a lake outlet, while in allochthonous material. Halesus spp. show a H. digitatus was found in most localities in the low coordinate score on the first axis resulting in subalpine zone. Apatania stigmatella showed a a lower coordinate score for zone 5 than expected. different distribution pattern than the other Apa tania species. Itwas not recorded at altitudes over The most obvious change in species composition 1220 m a.s.1. Solem (1985a) describedA. stigma according to the first axis occurred between zones 193 Reiso & Brittain: Distribution of Trichoptera in Jotunheimen Mountains, Norway 6 (1300-1350 m a.s.!) and 7 (1350-1400 m a.s.l). ptcra) i 0vre Heimdalsvatn, 0st-Jotunheimen (In According to 0sthagen & Egelie (1978), 1350 m Norwegian). 64 pp. Thesis, Zoological Museum, a.s.l is the transition between the low alpine and University of Oslo. the mid-alpine. This is probably due to the reduc Frost, S., Huni, A. & Kershaw, W. E. 1971. Evaluation ofa kicking technique for sampling stream bottom tion in riparian vegetation combined with lower fauna. Can. 1. Zoo!' 49,167-173. temperatures. Gislason, G. M. 1981. Distribution and habitat prefer ences of Icelandic Trichoptera. Pp 99-109 in Acknowledgements. We wish to thank Gunnar Hal Moretti, G.P. (ed.). Proceedings of the 3rd into vorsen for valuable comments on the manuscript, and Symp. on Trichoptera University ofPerugia Jul. 28 John Solem for important information on some ofthe - Aug. 2, 1980. Junk, The Hague, London. species. Research facilities in 0vre Heimdalen were Higler, L. W. G. & Solem, 1. O. (1986). Key to the provided by the Zoological Museum, University of larvae ofNorth-West European Potamophylax spe Oslo. The Norwegian Water Resources & Energy Di cies (Trichpotera, Limnephilidae) with notes on rectorate (NVE) kindly supplied temperature data from their biology. Aquatic Insects 8, 159-169. the outflow of 0vre Heimdalsvatn. Hiley, P., D. 1976. The identification of British limne philid larvae (Triehoptera). Syst. Ent. 1,147-167. REFERENCES Johannessen, T. W. 1978. The local climate ofthe 0vre Heimdalen valley. Holarc. Eco!. 1,93-102. Allan, D. J. 1995. Strcam Ecology. 388 pp. Chapman Larsson, P & Tangen, K. 1975. The input and the sig & Hall, Alden Press, Oxford. nificance of particulate terrestrial organic carbon Andersen, T. & Tysse, A. 1984. Lifc cycle of Chaeto in a SUbalpine freshwater ecosystem. Pp. 351-320 pteryx villosa (Fabricius, I 798)(Trichoptera: Lim in Wielgolaski, F. E. (ed.). Fennoscandian tundra nephilidae) in a lowland- and a Mountain- Stream ecosystems. Part 1. Springer, Berlin. in Western Norway. Aquatic Insects 6, 217-232. Larsson, P., Brittain, J.E., Lien, L., Lillehammer, A. & Andersen, T. & Wiberg-Larsen, P. 1987. Revised check Tangen, K. 1978. The lake ecosystem of 0vre list ofNW European Trichoptera. Ent. Scand. 18, Heimdalsvatn. Holarc. Ecol. I, 304-320. 165-187. Lillehammer, A. 1974. Norwegian stoneflies. II. Dis Andresen, T. 1979. Trichoptera. Fauna of Hardanger tribution and relationship to the environment. Norsk vidda 13, 1-18. ent. Tidsskr. 21, 195-250. Brittain, 1. E. 1978. Semivoltism in mountain popu Lillehammer, A. 1975. Norwegian stoneflies. III. Field lations of Nemurella pictetii (Plecoptera). Oikos studies on ecological factors inflU\;ncing distribu 30, 1-6. tion. Norw. J. Entomo\. 22, 71-80. Brittain, J. E. 1983. The influence of temperature on Lillehammer, A. 1984. Distribution, seasonal abundance nymphal growth rates in mountain stoneflies (Ple and emergence ofstoneflies (Plecoptera) in the 0vre coptera). Ecology 64,440-446. Heimdalen area of the Norwegian Jotunheimen Brittain, J. E. 1990. Life history strategies in Epheme Mountains. Fauna norv. Ser. B 31, 1-7. roptera and Plecoptera. Pp.I-12 in 1. C. Cambell Lillehammer, A. & Brittain, 1. E. 1978. The inverte (ed.) Mayflies and Stoneflies: Life Histories and brate fauna of the streams in 0vre Heimdalen. Biology. Kluwer Academic Publishers, Dordrecht. Holarc. Eco\. 1,271-274. Butler, M. G. 1984. Life histories of aquatic insects. Lillehammer, A., Brittain, 1. E., Saltveit, S. 1. & Nielsen, Pp. 24-55 in Resh, V. H. & Rosenberg, D. M. (eds.). P. S. 1989. Egg development, nymphal growth and Ecology ofaquatic insccts. Praeger Publishers, New life cycle strategies in Plecoptera. Holare. Eco!. 12, York. 173-186. Decamps, H. 1966. Sur la determination des femelles Malicky, H. 1983. Atlas of European Trichoptera. 298 de Potamophylax latipennis (Curt.) Neboiss et Pota pp. W. Junk, The Hague, London. mophylax cinglllatlls (Stephens) (Trichoptcra). Odinsson, A. 1. 1988. Research on Benthic Animal Life AnnIs Limnol. 2, 537-541. in the Blanda River System. University ofIceland, Edington, J. M & Hildrew, A. G. 1981. A key to to the Reykjavik. caseless caddis larvae ofthe British Isles, with notes Petersen, R. c., Gislason, G. M. & Vought, L. B.-M. on their ecology. Sci. Pub!. Freshwat. Bio!. Ass. 1995. Rivers ofthe Nordic countries. Pp.295-341 43,1-92. in Cushing, C. E., Cummins, K. W. & Minshall G. Eikeland, T. 1. 1982. Livssyklus og fodeopptak hos W. (eds.). River and stream ecosystems, Elsevier, Polycentropus flavomaculatus (Pictet) (Tricho Amsterdam. 194 Norw. J. Enlomol. 47, 185-195.2000 Reiso, S. 1999. Distribution and Ecology ofTrichoptera in a Norwegian Moutain area, 0vre Heimdalen. 49 pp. Thesis, Agricultural University ofNorway, As. Sand, K. 1997. Life cycle studies of Ephemeroptera and Plecoptera in a Norwegian mountain area, 0vre Heimdalen. 36 pp. Thesis, Zoological Museum, University of Oslo. Skjeseth, S. & Kloster, A. E. 1978. The geology of the 0vre Heimdalen valley. Holarc. Ecol. I, 89-92. Solem, J. O. & Gullefors, B. 1995. Trichoptera, cad disflies. Pp. 223-255 in NillsonA. (ed). The Aquatic Insects ofNorth Europe. Apollo Books, Stenstrup, Denmark. Solem, J. 0.1969. Livssyklus og biologi tilPhryganea bipunctata (Retzius) ogAgrypnia obsoleta (Hagen) (Trichoptera, Phryganeidae). Thesis, University of Trondheim, Trondheim. Solem, J. O. 1985a. Norwegian Apatania Kolenati (Trichoptera: Limnephilidae): identification oflar vae and aspects oftheir biology in a high- altitude zone. Ent. Scand. 16, 161-174. Solem, 1. O. 1985b. Distribution and biology of cad disflies (Trichoptera) in Dovrefjell mountains, Cen tral Norway. Fauna norv. Ser. B 32, 62-79. Ter Braak, C. J. F. 1995. Ordination. Pp. 91-173 in Jungman, R. H. G., Ter Braak, C. 1. F. & Van Ton geren, O. F. R. (eds.). Data analysis in community and landscape ecology. Cambridge University Press, Cambridge. 0sthagen, H. & Egelie, K. 1978. The vegetation of the 0vre Heimdalen valley. Holarc. Ecol. I, 103-106. Aagaard, K., Solem, J.O., Lillehammer, A., Hanssen, 0., Nost, T. & Dalen, T. 1989. Utbredelse, sonering og arsvariasjoner hos bunndyr i Atna og Atnasjoen. Miljovirkninger av vassdragsutbygging (In Norwe gian). MVU-rapportnr. B57: 21-26, NTNF, Trond heim. Received 13 July 2000, accepted 3 November 2000 195 Short communication The pteromalid subfamily Cerocephali trap). For further information about the localities, nae (Hymenoptera, Pteromalidae) in see Hanssen & Hansen (1998) for the former Norway record, and Hansen & Falck (2000) for the latter. Cerocephala rufa (Walker, 1833) Lars Ove Hansen VE Larvik: Middagskollen (EIS 19) Icr 15 June 6 July 1997, 19 6 July - 20 Aug. 1997 leg. A. Fjellberg (Malaise-trap). Hansen, L. O. 2000. The pteromalid subfamily Ceroce phalinae (Hymenoptera, Pteromalidae) in Norway. Biology and distribution Norw. J. Entomo!' 47, 196. All the known N European species ofCerocephali The pteromalid subfamily Cerocephalinae is recorded nae are parasitoids of beetle larvae. The biology of for the first time in Norway. Cerocephala cornigera C. cornigera is poorly known, but in Poland it has Westwood, 1832 is recorded from Oslo and Asker (AK) been hatched from the bark beetle Leperesinus orni and Cerocephala rufa (Walker, 1833) is recorded from (Col., Scolytinae) (Graham 1969). C. rufa may de Larvik (VE). Notes on distribution and biology are brief velop as a hyperparasitoid of certain wood-boring ly given. beetles of Scolytinae or Anobidae (Col.) via a pri mary parasitoid of Doryctinae (Hym., Braconidae) Key words: Cerocephala cornigera, Cerocephala ruja, (Gauld & Bolton 1988). Both species are recorded Cha1cidoidea, Pteromalidae, Cerocephalinae. from C Europe and Sweden, but C. cornigera is also recorded in Denmark (Graham 1969, Hansson 1991). Lars Ove Hansen, Zoological Museum, University of Oslo. Sarsgate 1, NO-0562 Oslo. Norway. Acknowledgements. I am indebted to Oddvar Hanssen for help during the field work, and to Arne Fjellberg who kindly donated insect material to the Zoological museum. Also thanks to 0stensjovannets Venner (i.e. Friends of Introduction 0stensjovannet), represented by Finn Gulbrandsen, who The family Pteromalidae includes fifteen sub financed the insect study at Lake 0stensjovannet. families in NW Europe (Graham 1969, Gauld & Bolton 1988). Among these, Cerocephalinae is cer References tainly the most peculiar, often with heavily Gauld, I. & Bolton, B. (eds). 1988. The Hymenoptera. tridentated heads, and with tufts of black bristles British Museum (Natural History). 332 pp. Oxford on each parastigmus. Five species from three gen University Press. era are recorded in NW Europe (Graham 1969, Graham, M.W.R. de V. 1969. The Pteromalidae of Gauld & Bolton 1988, Hansson 1991). However, Northwestern Europe (Hymenoptera: Chalcidoi dea). Bul!. Brit. Mus.(Nat. Hist.) Ent. supp!. 16, 1 this is the first record of this subfamily in Nor 908. way. The specimens are determined by the means Hansen, L.O. & Falck, M. 2000. Insektfaunaen ved ofthe keys in Graham (1969), and all material are 0stensjovannet, Oslo kommune. 75 pp. 0stensjo deposited in the collections at the Zoological mu vannets venner, Oslo. seum of Oslo. Hanssen, O. & Hansen, L.O. 1998. Verneverdige insekt habitater. Oslofjordomradet. NINA Oppdragsmel The records ding 546, 1-132. Hansson, e. 1991. ACatalogue ofthe Chalcidoidea (Hy Cerocephala cornigera Westwood, 1832 menoptera) described by e.G. Thomson, with a AK Asker: Bjorkas (EIS 28) 192 July - 24 Aug. checklist of Swedish species. Ent. Scand. Supp!. 38, 1-70. 1994 leg. L.O. Hansen & O. Hanssen (Malaise trap); Oslo: 0stensjovannet, Abildso (EIS 28) 1 Received 10 October 2000, 31 July 1996 10' 19, leg. Morten Falck (Malaise accepted 17 November 2000 196 Norw. J. Entomol. 47, 197-198. 2000 Platyderus ruficollis (Marsham) (Coleoptera, Carabidae) new to Scandinavia. John Skartveit, Frode 0degaard & Torstein Solh"y Skartveit, 1., 0degaard, F. & Solhey, T. 2000. Platyderus ruficollis (Marsharn) (Coleoptera, Carabidae) new to Scandinavia. Norw. 1. Entomo!. 47,197-198. The ground beetle Platyderus ruficollis (Marsham) is recorded as new to Scandinavia from a small islet near Arendal, South Norway. The nearest previous records of this eurytopical but rare species originate from the Netherlands and Britain. John Skartveit, Museum olZoology, Museplass 3. N-5007 Bergen, Norway Frode 0degaard, Norwegian Ins. for Nature Research, Tungasletta 2, N-7485 Trondheim, Norway Torstein Solhoy, Department ofZoology, Univ. ofBergen, Allegaten 36, N-5007 Bergen, Norway INTRODUCTION DESCRIPTION OF THE LOCALITY One of us (TS) collected a material of epigaeic The islet Svend Johnsens Holme is small, approxi arthropods using pitfall traps in a number ofsmall mately 100 by 150 meters, and sparsely vegetated, islets outside Arendal, South Norway in 1982 with heather (Calluna vulgaris) dominating on the 1983. At AAY Arcndal: Svend Johnsens Holme higher parts ofthe islands, and with low herbs and (EIS 6, UTM 32 VMK 837746), the material in grasses close to the seashore and in several crev cluded five specimens of the carabid Platyderus ices. The beetle fauna was dominated by xerophi ruficollis (Marsham). This is the first North Euro lous and heliophilous species and was rather rich pean record of the genus Platyderus. As this un in carabids, with 21 species collected (Skartveit expected species is not described in the identifi & Solhoy, unpublished observations). cation literature commonly used for Scandinavian Platyderus species are considered to be mostly carabids, it is briefly described and illustrated here. montane in South and Central Europe (Trautner & Geigenmiiller 1987), whereas the Norwegian BRIEF DESCRIPTION OF THE SPECIES locality is coastal. Aukema & Baars (1986), how A medium-sized, rather slender carabid, 5-7 mm ever, found the species in small numbers on a long, head dark brown to black, body and append heathland on the island Texcl in the Netherlands. ages reddish-brown. Its habitus (Figure 1) most Platyderus ruflcollis is an eurytopical species, oc resembles that of Synuchus vivalis (Illiger), but curring in forests as well as in open country (Koch may also look like a small Pterostichus or Olistho 1989), but despite this it is quite rare. Its distribu pus species. The pronotum is relatively large and tion is South-Western Europe, being hitherto re rounded, with the basal foveae deep and prolonged corded from Ireland, England, the Netherlands, forward as sharp furrows almost reaching to the France, Switzerland, Spain, Portugal and Italy middle of the pronotum. The epipleurae are not (Aukema and Baars 1986). crossed as they are in Pterostichus, and the elytra each have two small dorsal punctures in the third POSSIBLE ORIGIN OF THE POPULATION stria. The mentum carries a median tooth like in The population is rather remote from any previ Agonum, and the claws are smooth. ously known populations of this species. The 197 I Skartveit, 0degaard & Solhew: Platyderus ruficollis (Coleoptera, Carabidae) new to Scandinavia rujicol/is should be included as a vulnerable (V) 1~:::~:~;;~~:e:~~:~;it~n:::t;~eOra~:il~~~~ species in the next Red list for Norwegian beet :century. Numerous sailships landed in Norway les. (rom various parts of the world in this area. The ,islet is situated at the mouth of the river Nidelva References land would be a quite likely anchoring site for sail Aukcma, B. & M.A. Baars 1986. Platyderus n~ficollis Ivessels. In particular, many ships coming for tim (Marsham, 1802), een nieuve Nederlandse loop ber entered from England and the Netherlands, and kcvcr van Texel (Coleoptera, Carabidae). Entomo may have dumped earth or sand used as ballast. logische Berichten (Amsterdam) 46, 19-20. Any beetles in the ballast material may have floa Koch, K.C. 1989. Die Kiifer Mitteleuropas. Okologie ted ashore. Since the islet lies at the mouth of a I. 440 pp. Goecke and Evers, Krefeld. rather large river, the surface water is quite fresh Lindroth, C.H. 1985-1986. The Carabidae (Coleoptera) and beetles may survive in it for some time. Intro ofFennoscandia andDenmark. Fauna entomologica Scandinavica IS. 497 pp. E.l. Brill/ Scandinavian duction of this species with the sailship trade is Science Press Ltd, Leidenl Copenhagen. thus plausible. Trautner, 1. & K. Geigenmuller 1987. Tiger beetles ground beetles. l1lustrated key to the Cicindelidae REDLIST STATUS - SUGGESTION and Carabidae of Europe. 488 pp. 10sef Margraf, publisher, Aichtal, Gennany. The species is currently known from only one population in Norway. The locality is not protected Received 4 September 2000, (Arild Pfaff, personal comm.), and lies in area accepted 25 September 2000 where nature is under heavy pressure from rec reational activities. We suggest that Platyderus Imm Figure 1. Platyderus rufical/is (Marsham), habitus. 198 Norw. J. Entomo/. Contents: Vol. 46-47 Norwegian Journal of Entomology Volume 46 • 1999 Articles Greve, L. Notes on tephritid flies (Diptera, Tephritidae) from Norway 1 Aarvik, L. & Bakke, A. The moth (Lepidoptera) fauna of coastal spruce forest at Almedalen Forest Reserve, Namsos, and continental forest in Lierne, Central Norway 9 Kvamme, T. Notes on Norwegian ants (Hymenoptera, Formicidae) 19 Hagvar, S. Saproxylic beetles visiting living sporocarps of Fomitopsis pinicola and Fomes fomentarius 25 Falck, M. Lejogaster tarsata (Megerle in Meigen, 1822) (Diptera, Syrphidae) new to Norway .. 33 S~mme, L. & Birkemoe, T. Demography and population densities of Folsomia quadrioculata (Collembola, Isotomidae) on Spitsbergen 35 Hansen, L.a. & Greve, L. New records of Triozidae (Hem., Psylloidea) from Norway 47 Hauge, E. & Thingstad, PG. Spiders (Araneae) from the Kvitbergvatnet area in Saltdal, northern Norway 49 Hansen, L.a. & Berggren, K. The genus Nothochrysa (Planipennia, Chrysopidae) in Norway 57 Hagvar, S. New data on the distribution ofNorwegian Hemiptera Heteroptera 61 0degaard, F. Invasive beetle species (Coleoptera) associated with compost heaps in the Nordic countries 67 Ekrem, T. Tanytarsus mancospinosus sp. n. (Diptera, Chironomidae) from eutrophic lakes in Europe 79 Falck, M. & Greve, L. The distribution ofbee flies (Diptera, Bombyliidae), except the genus Villa, in Norway 89 Short communications Greve, L. Gampsocera numerata (Heeger, 1858) (Diptera, Chloropidae) new to Norway 8 Fjellstad, B.M. Hydraecia ultima (Holst, 1965) (Lep., Noctuidae) recorded in Norway 18 Skartveit, J. Adistemia watsoni (Wollaston) (Col., Latridiidae) recorded from Norway 46 Andersen, T. Moth flies (Diptera, Psychodidae) from the Faroes 66 Olsen, TJ. Myrmica mgulosa Nylander, 1849 (Hymenoptera, Formicidae) new to Norway 88 Olsen, TJ. New records of Psylloidea and Auchenorrhyncha (Homoptera) from Norway 110 Book reviews • Bokanmeldelser 24 199 Contents 1999--2000 Norwegian Journal ofEntomology Volume 47 • 2000 Articles Kobro, S., Teksdal, A.E. & Andersen, A. Cereals as host plants for thrips (Thysanoptera) in Norway I 0degaard, F. & Ligaard, S. Contribution to the knowledge of Norwegian Coleoptera 7 Fjellberg, A. & Jucevica, E. A new species ofArchisotoma Linnaniemi, 1912 from the Baltic coast of Latvia (Collembola, Isotomidae) 21 Gogstad, G.O. Acid rain and the disappearance of the apollo butterfly (Parnassius apollo (L., 1758)) from coastal areas in Norway 25 Andersen, J., Olberg, S. & Haugen, L. Saproxylic beetles (Coleoptera) ofTroms and western Finnmark, northern Norway with exceptional distribution in Fennoscandia 29 Coulson, SJ. A review of the terrestrial and freshwater invertebrate fauna of the High Arctic archipelago of Svalbard 41 Haenni, J.-P. & Greve, L. New records of Norwegian Scatopsidae (Diptera) 65 Hauge, H. & Hansen, L.O. New records of spiders (Araneae) from the Oslofjord area, SE Norway 73 Aakra, K. New records of spiders (Araneae) from Norway with notes on epigynal characters of Philodromus fuscomarginatus (De Geer) (Philodromidae) and Araneus sturmi (Hahn) (Araneidae) 77 Greve, L. New records of Norwegian Lauxaniidae (Diptera) 89 Aakra, K. Ag)'neta mossica (Schikora, 1993) (Araneae, Linyphiidae) in Norway 95 Johansen, M., Elliott, J.M. & Klemetsen, A. Diel fluctuations of invertebrate drift in a Norwegian stream north of the Arctic Circle 101 Reiso, R. & Brittain, J.E. Life cycle, diet and habitat of Polycentropus flavomaculatus, Plectrocnemia conspersa and Rhyacophila nubila (Trichoptera) in 0vre Heimdalen, Jotunheimen Mountains, Norway... 113 Tommeras, B.A. Individual variation in behavioural attraction and corresponding olfactory receptor capability to enantiomers of the pheromone ipsdienol in the pine engraver Ips pini 125 Stur, E. & Andersen, T. A new species ofPhysoneura Ferrington et S 200 Norw. J. Entomol. Contents: Vol. 46-47 Short communications Olsen, K.M. Mitostoma chrysomelas (Hennann, 1804) (Opilionida, Nemastomatidae), a harvestman new to Norway 24 Olsen, K.M. Haplophilus subterraneus (Shaw, 1789), a centipede (Chilopoda, Geophilomorpha) new to Norway , , 63 Greve, L. & Johanson, K.A. Malacomyia sciomyzina (Haliday, 1833) (Diptera, Coelopidae) recorded from Norway , , 72 Hansen, L.a. The family Signiphoridae (Hymenoptera, Chalcidoidea) in Norway 76 Hansen, L.a. Euides speciosa (Boheman, 1845) (Homoptera, De1phaeidae) in Norway 148 Greve, L. The genus Pteromicra Lioy, 1864 (Diptera, Sciomyzidae) in Norway 182 Hansen, L.a. The pteromalid subfamily Cerocephalinae (Hymenoptera, Pteromalidae) in Norway 196 Book reviews· Bokanmeldelser 19, 183 201 Instructions to authors Instructions to authors with extended information on electronic submittence Manuscripts authors, it should be referred to as Smith et al. (1990). Multiple references in the text are given The language is English or occasionally Norwe in chronological orders (Smith 1975, Green 1980, gian with an cxtended English summary. Black & White 1998). All references (but not any Manuscripts, double spaced on one side of thc that has not been cited in the text) should be listed paper must be submitted in duplicate. Separate in alphabetical order at the end of the paper. 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Examples: do not contain halftones (photographs or scans). Male(s): I (J = I M 13 (J(J = 13 MM In these figures you may «shade» different regions Female(s): I 9 = 1 F 1699 = 16 FF by using line shading or hatching, or coarse tints Worker(s): 1 9 = 1 V 1899= l8VV ofgrey. Note that black text should be used over Hybrid(s): 1 9 = I H 12 gg = 12 HH light to mid-greys, and white text over dark grey or black shades. Figures Individual illustrations should be exported or saved Figures must be numbered consecutively and all as EPS files and given an EPS extension (e.g. figures must be referred to in the text. Write «Fig Figl.eps). Filenames should not have more than 8 ure» in full. Names(s) of the author(s) should be characters with 3 extension letters. These figures written in the margin of each figure. The size of should be saved in 1 bit, unless they contain tints, the figure must not exceed 210 x 290 mm (stand in which case they should be saved in 8 bit. ard A4). The author should take into considera If possible, all fonts should be embedded in the tion that most figures have to be reduced. Please figures and, ifnot, a note should be made ofnon take in considerations that the text block is 142 x embedded fonts. Iffont embedding is unavailable 205 mm, and the column with is 68.5 mm. In line please use standard system fonts (ideally Arial or drawings the line thickness should not be less than Helvetica). Font size should be appropriate to the 0.25 mm after reduction, and capital letters not final product, an indication is that a figure at its smaller than 2.0 mm. Choose contrasting patterns final size should have 10 or 12 pt text. and avoid fine tone rasters. Maps and morpho logical illustrations, e.g. pictures ofinsects, should Please try to ensure that a PostScript printer driver include a scale bar. is used to generate the EPS file. An Adobe EPS driver is recommended. Most art programs give the option to «save as» EPS or TIFF, or «export» the file in EPS or TIFF We also accept TIFF files which are cropped close format. These two formats are far more compat to the edge of the figure to minimize the white ible with image-editing software and supported space surrounding the image. The TIFF should by more programs than program-specific file for be saved at a resolution of800 d.p.i. (dots per inch) mats. We do not accept figures as program-files at final size. Supplying uncompressed TIFFs is (e.g. Excel, Power-point). Please use the follow preferable but ifthe image size is very large com ing file formats: pression software can be used. We would then like to know the type of compression used (i.e. LZW, EPS. The Encapsulated PostScript (EPS) language WinZip, etc.). For figures consisting ofmore than file format can contain both «vector-graphics» (i.e. one element (e.g. parts (a), (b) and (c)), please the image is described as lines, curves and letter supply the different parts separately (i.e. (a) sup ing in specific fonts) and «bitmap-graphics» (i.e. plied in a different file to (b)). the image is made up ofpixels) and is supported by virtually all graphic, illustration and desktop Line drawings should be scanned at an original publishing programs. resolution that will permit a final output ofat least 800 d.p.i. (dots per inch). If the figure is to be TIFF. The Tagged-Image File Format (TIFF) is reduced to 50% of the original size the scanning used to exchange files between programs and op- 203 Instructions to authors resolution should be at least 400 d.p.i. It would plied in a different file to (b)). An Adobe EPS jbe most helpful if you could state if reduction is driver is recommended. Inecessary. Always enclose high quality paper-copies ofyour IAlways enclose high quality paper-copies ofyour photograph(s) / black-and-white halftone figure(s) figure(s) in case we are not able to use the elctronic in case we are not able to use the elctronic ver version(s). sion(s). Photographs / black-and-white halftones. These must be in black and white and of high quality Proofs and reprints with good contrasts. It is important that these fig The author will receive one set of proofs which ures are supplied to the correct resolution as oth must be returned without delay. Extensive altera erwise we will not be able to use them. For print tions will be charged to the authors. An order form ing purposes halftone figures need to have a mini for reprints must be returned with the proofs. The mum resolution of 300 d.p.i. (dots per inch) at journal does not provide free reprints. final size, otherwise the image will be blurred Ifyou have any queries please contact the editorial when printed. Computer screens display a reso secretary: Lars Ove Hansen. lution of only 72 or 96 d.p.i. so it is important to E-mail: [email protected] check that the halftone has a high-enough resolu tion for printing purposes. The minimum shade ofgrey should ideally be 20% (otherwise the minimum greys will reproduce as extremely faint when printed) and the darkest greys should be no greater than 80% (otherwise they will show as black when printed). Try and make the shades as coarse and distinct as possi ble for maximum clarity. Individual halftones should be exported or saved as TIFF files and given a TIFF extension (e.g. Figl.tif). Filenames should not have more than 8 characters with 3 extension letters. These figures should be saved in 8 bit. We prefer TIfF files saved close to the extremity of the figure to minimize the white space surrounding the image. Supply ing uncompressed TIFFs is preferable but if the image size is very large compression software can be used. We would like to know the type ofcom pression used (i.e. LZW, WinZip, etc.). If you cannot supply figures in TIFF format, we can also accept EPS files of your grey-scales. They should be exported or saved in 8 bit, with an EPS extension (e.g. l-l.eps). Filenames should not have more than 8 characters with 3 extension letters. Each file should be centred on the page and saved at final size with the correct orientation. For figures consisting of more than one clement (e.g. parts (a), (b) and (c)) Please supply the different parts separately (i.e. (a) sup 204 INSTRUCTIONS TO AUTHORS Green 1980, Black & White 1998). All references (but not any that has not been cited in the text) should be The language is English or occasionally Norwegian listed in alphabetical order at the end of the paper. In with an extended English summary. English reference lists, 0 is equal to 0 and Ais equal to Aa. Names ofjournals are abbreviated according to in Manuscripts, double spaced on one side of the paper ternational standards, e.g. as in BIOSIS (Biological must be submitted in duplicate. Separate sheets should Abstracts). be used for (I) title page with authors names, (2) ab stract followed by the name(s) and postal address(es) Examples: of the author(s), (3) tables, (4) numbered figures and (5) legends to figures. Acknowledgements should be Journal papers gathered under a single heading at the end of the text. Chant, D.A. & McMurtry, J.A. 1994. A review of the All manuscripts will be considered by referees before subfamilies Phytoseiinae and Typhlodrominae acceptance. (Acari: Phytoseiidae). Int. 1. Acarol. 20, 223-310. Abstract should not exceed 300 words and should cover Book the main results and conclusions of the paper. A list of Borror, 0.1., Tripletorn, c.A. & Johnson, N.F. 1989. up to five key words may be added on a separate line An introduction to the study of insects. Sixth edi below the abstract. tion. 875 pp. Saunders College Publishing, Phila delphia Tables are typed on separate sheets and numbered con secutively. Each table must have a heading. Write «Ta Chapter in book ble» in full both in the text and table heading. Avoid Dennis, R.L.H. & Williams, WR. 1995. Implications vertical lines in the table. of biogeographical structures for the conservation of European butterflies. Pp. 213-230 in Pullin, A.S. Figures must be numbered consecutively and all fig (ed.), Ecology and conservation of butterflies. ures must be referred to in the text. Write «Figure» in Chapman & Hall, London. full. Names(s) of the author(s) should be written in the margin of each figure. The size of the figure must not Computer files. After acceptance the author(s) will be exceed 210 x 290 mm (standard A4). The author should asked to submit a computer disc (3.5") or a compact take into consideration that most figures have to be re disc (CD-rom) containing the final version of the paper duced. In line drawings the line thickness should not be along with the revised manuscript itself. We may also less than 0.25 mm after reduction, and capital letters accept Attachments to e-mail, but only in agreement not smaller than 2.0 mm. Choose contrasting patterns with the editorial secretary. Operating systems and word and avoid fine tone rasters. Photographs must be in processing program must be written on the disc. We black and white and of high quality with good contrasts. recommend that the text is saved as RTF «Rich text Maps and morphological illustrations, e.g. pictures of format». Always separate text and figures in the elec insects, should include a scale bar. tronic version. Further details will be supplied by the editor. Nomenclature. Scientific names of genera and species should be written in italics. The name of the author and Proofs and reprints. The author will receive one set of year of description (if appropriate), separated by a proofs which must be returned without delay. Exten comma, should be included the first time the name of sive alterations will be charged to the authors. An order an insect or another terrestrial arthropod is mentioned form for reprints must be returned with the proofs. The in the text, e.g. Rhyacophila nubila (Zetterstedt, 1840). journal does not provide free reprints. Names of authors should be written in full. LocaJities. In faunistic papers the names of Norwegian localities should be according to K.A. 0kland (1981), Fauna (Oslo) 34, 167-178, and preferably the EIS number should be added in brackets. References. Citations in the text should be written as Black (1992), (White 1995) or (Black & White 1998). When a paper has more than two authors, it should be referred to as Smith et al. (1990). Multiple references in the text are given in chronological orders (Smith 1975, CONTENTS Norw. J. Entomol. 47 (2) 2000 ISSN 1501-8415. Articles Johansen, M., Elliott, lM. & Klemetsen, A. Diel fluctuations of invertebrate drift in a Norwegian stream north of the Arctic Circle 101 Reiso, R. & Brittain, J.E. Life cycle, diet and habitat of Po(vcentropusjlavomaculatlls, Plectrocnemia conspersa and Rhyacophila flubila (Trichoptera) in 0vre Heimdalen, Jotunheimen Mountains, Norway... 113 Tommen'is, B.A. Individual variation in behavioural attraction and corresponding olfactory receptor capability to enantiomers of the pheromone ipsdienol in the pine engraver Ips pini 125 Stur, E. & Andersen, T. A new species of Ph)'soneura Ferrington et Srether, 1995, from Ecuador (Chironomidae, Orthocladiinae) 131 Martinsen, L. & Soli, G. E. E. Description of females of three species of Ectrepesthoneura Enderlein (Diptera, Mycetophilidae) 137 Aakra, K., Bretten, A. & Frengen, O. Spiders (Araneae) new to Norway 149 Aakra, K. Noteworthy records of spiders (Araneae) from centra) regions of Norway 153 Riedel, M., Hansen, L.O. & Berg, 0. Ichneumonidae (Hymenoptera) new for the fauna of NOlway 163 Berg, 0. Aculeata of Norway. 3. Eleven species of bees new to Norway (Hymenoptera: Apoidea) 177 Reiso, R. & Brittain, lE. Life cycle, diet and habitat of Polycentropllsjlavomaculatus, Plectrocllemia cOllspersa and Rhyacophila nubila (Trichoptera) in 0vre Heimdalen, Jotunheimen Mountains, Norway... 185 Skartveit, J., 0degaard, F. & Solhoy, T. Platyderus rujicollis (Marsham) (Coleoptera, Carabidae) new to Scandinavia 197 Short communications Hansen, L.O. Euides speciosa (Boheman, 1845) (Homoptera, Delphacidae) in Norway 148 Greve, L. The genus Pteromicra Lioy, 1864 (Diptera, Sciomyzidae) in Norway 182 Hansen, L.O. The pteromalid subfamily Cerocephalinae (Hymenoptera, Pteromalidae) in Norway 196 Book reviews· Bokanmeldelser 183 The present journal is a continuation of Fauna norvegica Series B (Norwegian Journal of Entomol ogy). With Volume 45 (1998) the publication of this journal by the Foundation for Nature Research and Cultural Heritage Research (NINA-NIKU) has been discontinued. Starting with Volume 46, the journal is published as Norwegian Journal of Entomology by the Norwegian Entomological Society. Editor is Professor La:uritz S0mme, University of Oslo.