Ent. Tidskr. 138 (2017) fauna on Stora Karlsö Snapshot of the Hymenopteran fauna of Stora Karlsö

JULIA STIGENBERG, JOSEF BERGER, MATTIAS FORSHAGE, NIKLAS JOHANSSON, AR- TUR LARSSON, OLE LØNNVE, ALEXEY RESHCHIKOV, HEGE VÅRDAL & IKA ÖSTER- BLAD

Stigenberg, ., Berger, J., Forshage, ., Johansson, ., Larsson, A., Lønnve, ., Resh- chikov, A., Vårdal, . & Österblad, I.: Snapshot of the Hymenopteran fauna of Stora Karl- sö. [Ögonblicksbild av stekelfaunan på Stora Karlsö.] – Entomologisk Tidskrift 138 (1): 71-91. Uppsala, Sweden 2017. ISSN 0013-886x.

Stora Karlsö is a small island close to Gotland in the Baltic Sea of which the Hymenopteran fauna has not been extensively studied before. In August 2014, a team of eight persons carried out an inventory of Hymenoptera, mainly the parasitoid and , on the island. Sampling was done with Malaise traps for a period of 22 days, complemented with vegetation sweeping, branch shaking and opportunistic handpicking during a five day sojourn. As a result, about 200 species of parasitoid wasps and 14 sawflies are reported for Stora Karlsö for the first time. Eleven species are reported as new to Sweden: The cornubiae Benson, 1931, the gasteruptiid opacum (Tournier, 1877), the diapriid Spilomicrus rufitarsis(Kieffer, 1911), the eulophid Entedonomphale bulgarica Boyadzhiev & Triapitsyn, 2007, the braconids Bracon rozneri Papp, 1998 and Gnampto- don decoris (Förster, 1862), and the ichneumonids Bathythrix maculata (Hellén, 1957), Heterischnus filiformis (Gravenhorst, 1829), Lissonota picticoxis Schmiedeknecht, 1900, Mesochorus tipularius Gravenhorst, 1829, Ophion brevicornis Morley, 1915, and Plecto- chorus iwatensis (Uchida, 1928). Also the gasteruptiid Gasteruption opacum (Tournier, 1877) is reported new to Sweden based on a record from inventory by in 2013. This demonstrates how the knowledge of Swedish biodiversity can be substantially augmented by a short and intensive collecting expedition. We strongly recommend that other places in the country be subjected to similar efforts.

Julia Stigenberg, Mattias Forshage, Hege Vårdal, Naturhistoriska riksmuseet, Box 50007, 10405 Stockholm. -mail: [email protected], [email protected], [email protected]. Josef Berger, Department of Biology, Lund University, Lund. E-mail: [email protected] Niklas Johansson, Aspåsen/Baskarp, 56692 Habo E-mail: [email protected] Artur Larsson, ArtDatabanken, Box 7007, 75007 Uppsala. E-mail: [email protected] Ole Lønnve, BioFokus, Gaustadalléen 21, NO-0349 Oslo, Norway. E-mail: [email protected] Alexey Reshchikov, Sun Yat-sen University, 135 Xingangxi St. Guangzhou, 510275, China. Ika Österblad, [email protected]

Stora Karlsö (Fig. 1) is an island in the Baltic the west, north and northeast. The highest point Sea, belonging to Gotland both in administra- of the island is 50 m above sea-level. The island tive terms (Gotlands län & kommun) and as a consists of ca 400 million year old (Silurian) fos- faunal province. It is situated about 6 km off the sil coral reefs surrounded by layered limestone west coast of Gotland, has a subrectangular out- (Hedgren 2005, Johansson 2013). This exposed line and an area of about 2.5 km². A dominant and open alvar landscape represents a habitat feature of the landscape is a horse-shoe shaped which can be found almost exclusively in Swe- plateau forming steep cliffs along the coastline to den, on the islands of Gotland and Öland, and in 71 Julia Stigenberg et al. Ent. Tidskr. 138 (2017)

Figure 1. Map of Stora Karlsö by Stellan Hedgren. The Malaise trap locations are marked M1-M6. Karta över Stora Karlsö av Stellan Hedgren. Placeringen av Malaisefällorna är markerade M1-M6.

Estonia, with minor occurrences in southwest- the open areas which were once heavily grazed ern Finland (Eriksson & Rosén 2008). (Johansson 2013). For centuries, grazing kept the vegetation on A dominating shrub is the St. Lucie cherry Stora Karlsö low. Together with the geologi- (Prunus mahaleb) which is introduced and cal features, this gave rise to a unique flora and competes with the native blackthorn (Prunus fauna. When Linnaeus visited the island in July spinosa) (Johansson 2013, pers. obs.). Humans 1741, he reported that there was only one tree have been visiting the island since the stone age (an ash, which is still standing on the island) and (Hedgren 2005), and they have introduced many remarked that the grazing sheep were growing plant species (including walnut Juglans regia fat on the island (Linnaeus 1745). In 1887 graz- and sycamore Acer pseudoplatanus, though no ing was discontinued, resulting in a dramatic one as striking as the St. Lucie cherry (Broqvist change in the plant cover, with junipers (Juni- & Magntorn 1989, Johansson 2013). perus communis), pines (Pinus sylvestris) and More than for its vegetation, the island is deciduous trees spreading. In the mid-1990’, famous for the auk colonies in the cliffs (Guil- however, the island management launched an lemot Uria aalge and Razorbill Alca torda), effort to re-create conditions favourable for or- the most important ones in the entire Baltic. In ganisms dependent on grazing (Hedgren 2005). 1970, a nature reserve was created comprising The scree areas of the island are now covered the island and its surrounding water area within by deciduous forest dominated by ash (Fraxinus 1 km distance from the shoreline. It is consid- excelsior), Norway maple (Acer platanoides), ered one of the oldest nature protection areas in elm (Ulmus glabra and U. minor), rowan (Sor- the world, because its fauna and flora have been bus aucuparia) and Swedish whitebeam (Sorbus under effective protection since 1887 when the intermedia), with ivy (Hedera helix) climbing Karlsö Association for Hunting and Conserva- the trunks and cliffs. Juniper bushes that were tion (Karlsö Jagt- och Naturskyddsförening) rare in the late 1800s can now be found on the took over management of the island (http:// entire island and form shrubberies in many of www.storakarlso.se). 72 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö Hymenoptera isolation from similar habitat patches and their Parasitic wasps constitute a large part of ter- size, as these can potentially contribute coloniz- restrial diversity everywhere, while at ers (MacArthur & Wilson 1967, Hanski 1999). the same time being poorly known and usually Highly specialised species may have difficul- neglected in inventories and conservation man- ties maintaining viable populations on islands, agement. In Sweden, they comprise roughly a as their population dynamics inevitably reflect quarter of the animal species for which the in- host population fluctuations. Within the formation is so scant that the Swedish popula- community, such specialists are most promi- tions cannot be judged according to the IUCN nently represented by mono- and oligophagous red-listing criteria (Gärdenfors 2000). Recent herbivores and parasitoids. Parasitoids are par- activities within the framework of the Swedish ticularly vulnerable because of their position at Initiative (Karlsson et al. 2005) have a high trophic level. Comparing species richness started to clarify the diversity for some groups across islands in the archipelago of southwest- of parasitic wasps within the country, whereas ern Finland, Roslin et al. (2014) found that the other groups still remain very poorly known. species-area relationship was more pronounced However, it is widely acknowledged that para- in higher trophic levels: in the sequence go- sitoid wasps play a highly important role in the ing from plants through herbivores to primary ecosystem because by ultimately killing their and secondary parasitoids, species number de- host, they contribute to population regulation of creased successively more rapidly as a function other . Also, parasitoids are recom- of shrinking area. mended as indicators in applied conservation: According to Fukami (2015), community as- Shaw & Hochberg (2001) suggest that the oc- sembly – through sequential, repeated immigra- currence of a rare species is more valuable if it, tion from the regional species pool – is influ- on a particular location, acts as host of a spe- enced on the one hand by processes for which cialised parasitoid. Unfortunately, due to lack deterministic patterns can be postulated, such as of data, protective measures for these wasps are functional traits and phylogenetic structure, and rarely taken. Most information about the biolo- on the other by the rather more random timing gy of parasitoid wasps stems from agronomy re- and order of events. Influential events may be search on potential biocontrol agents, which for abiotic, such as flood or fire, or biotic, e.. colo- obvious reasons focuses on the most abundant nization by a species which positively or nega- species. Aculeate wasps are far better studied, tively affects the outcome of immigration by and thus well-known to be very relevant to con- another species. A species can exhibit unusual servation concerns particularly since the general abundance in the absence of otherwise signifi- public has become aware of the recent decline of cant predators, parasitoids, parasites or, perhaps honeybees. Sawflies are somewhat intermediate most importantly, when released from the in- in terms of knowledge, and mainly wood-living hibitory effect of competitors. If the organism species have been taken into consideration from communities of Stora Karlsö have ever reached a conservation viewpoint. stable states during the 12.000 years since its emergence after the last ice age, with severe ex- Island Biodiversity posure to the brute forces of wind and waves The number of species tends to increase with (Larje 2008), time after time these communi- area (Arrhenius 1921); this relationship has ties must have reassembled owing to changes been repeatedly observed and extensively used brought on by dramatic fluctuations in level and in ecological modelling (With 2016). Small salinity of the surrounding water, and not least habitat patches tend to hold small populations by human activities during 9.500 years. which are more likely to go extinct than are larg- populations, be it due to stochastic processes Previous inventories including Hymenoptera or susceptibility to changes in habitat quality. In spite of some scattered records, like ants col- Species composition and turnover are affected lected by the paleontologist Gustaf Lindström not only by habitat area but also by the degree of (Stolpe 1882), the entomological exploration of 73 Julia Stigenberg et al. Ent. Tidskr. 138 (2017)

Figure 2. The participating Hyme- noptera experts at the collecting effort in 2014. From left: Niklas Johansson, Artur Larsson, Mat- tias Forshage, Hege Vårdal, Julia Stigenberg, Ika Österblad, Alexey Reshchikov, Josef Berger. De deltagande stekelexperterna vid insamlingstillfället 2014. Från vänster: Niklas Johansson, Artur Larsson, Mattias Forshage, Hege Vårdal, Julia Stigenberg, Ika Österblad, Alexey Reshchikov, Josef Berger.

Stora Karlsö was far from methodical until some methods of collecting and observing organisms major collecting efforts by Hans Lohmander in are used (e.g. Laforest et al. 2013). The aim of 1927, by Olov Lundblad in 1950 and 1956, and this concentrated collection effort was to find especially by Nils Linnman continuously from and identify as many hymenopteran species as the 1930s on (Linnman 1965). Linnman was possible, focusing on the little known parasitic however mainly focused on Coleoptera, and wasps and sawflies. of these early visits the Hymenoptera material Our collecting expedition was approved by seems to have been very limited and mainly con- Länsstyrelsen Gotland (permit nr: Dnr 521-26- sisting of Lundblad’s aculeates. 14). In the 1980s, this exploration had a bit of a re- naissance with groups of entomologists spend- Material and methods ing time on the island most summers (Struwe Our inventory in 2014 was mainly carried out 1989). During these years, Lars Norén did a using Malaise traps (running 8-29 August) (Fig. fairly thorough inventory of major parts of the 3, 4). During five days (25-29 August) trap- aculeate fauna, as did Olle Högmo with the ants ping was supplemented with sweeping, branch- (Högmo 1988), and Carl-Cedric Coulianos with shaking and visual search (including scouring gall-making wasps, both gall wasps proper and windows). In addition, galls from Rosa sp. and gall-making sawflies (Coulianos 2010). Quercus robur were collected, but unfortunately In recent years, several hymenopterists have rearing was unsuccessful. During the inventory been visiting the island again, culminating in the week, the weather conditions were mostly fa- present inventory made by our team of eight ex- vourably dry and sunny, although the total sam- perts (Fig 2). Materials from most of these col- pling result presumably was somewhat nega- lecting events have been included in this paper, tively affected by strong winds. and are described under Material and methods The Malaise trap, a tent-like construction for below. collection of flying , has proved to be an We thought it would be interesting to address efficient method for collecting wasps, flies and such an extraordinary island locality with a midges in particular (Malaise 1937, Karlsson et methodology similar to that of a bioblitz where, al. 2005). In the present inventory, five Townes- during a short period of time, several different model Malaise traps were used, placed in differ- 74 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö

Figure 3. Julia emptying Malaise trap nr 2. In this trap we col- lected the Lissonota picticoxis Schmiedeknecht, 1900. Julia tömmer Malaisefälla nr 2. I denna fälla fann vi Lissonota picticoxis Schmiedeknecht, 1900. . ent habitats (Fig. 1). When the catch for the time go lanceolata, Polygala vulgaris, Scabiosa co- interval August 8-24 was inspected, the number lumbaria, Thalictrum sp., Veronica chamaedrys, of specimens at one location (M5) turned out to Vincetoxicum hirundinaria). be very low, and this trap was relocated for the Malaise trap #3, 57.27919°N 17.97448°E, remaining few days in order to maximize sam- elevation 11 m. Exposed, vegetated coastal pling efficiency (M6). Trap material was thus stone shore facing south. Alternating patches of collected from six strategically selected locali- vegetation and bare stone field. Trap placement ties. GPS coordinates are presented using WGS was at the border between open stone field and 84 system. The listed plants indicate the domi- deciduous shrubbery dominated by Prunus. (?) nating plants within 2 m from the trap? Anchusa arvensis, Anthriscus sylvestris, Cala- Malaise trap #1, 57.28821°N 17.97139°E, magrostis sp., Cirsium vulgare, Galium verum, elevation 5 m. Calcareous low herb pasture, be- Geranium robertianum, Geum urbanum, Plan- tween a few junipers and roses in a slope facing tago lanceolata, Potentilla argentea, Prunus south. Odontites ?vulgaris, Scabiosa columbar- mahaleb, Satureja acinos, Scabiosa columbar- ia, Thalictrum sp., Avenula pratensis/pubescens. ia, Sedum album, Sorbus aucuparia, Vincetoxi- Prunus mahaleb, Vincetoxicum hirundinaria, cum hirundinaria. Rosa sp., Juniperus communis, Fraxinus excel- Malaise trap #4, 57.28200°N 17.96597°E, el- sior, Fragaria viridis, Filipendula vulgaris. evation 24 m (Fig. 4). Moist calcareous meadow Malaise trap #2, 57.28730°N 17.97753°E, northwest of a less than 2 m tall rock face; the elevation 35 m (Fig. 3). Calcareous low herb trap placed adjacent to this. Along the rock face pasture, between sparse pine groves in a slope grew deciduous trees and shrubs (Crataegus facing west. There were a few junipers, small sp., Fraxinus excelsior, Juniperus communis, rowan saplings (Sorbus aucuparia) and Rosa sp, Prunus mahaleb), further from it there were but the locality was open, the ground covered by open meadow patches with Dactylis glomerata, mosses and lichens (Cladonia sp, Hypnum sp.), Festuca rubra, Filipendula vulgaris, Fragaria sparse grass (mainly Helictotrichon pratense viridis, Galium verum, Geranium robertianum, and Phleum phleoides) and low herbs (Achillea Hedera helix, Helictotrichon pratense, Hepat- millefolium, Filipendula vulgaris, Fragaria viri- ica nobilis, Hypericum perforatum, Odontites dis, Galium boreale, G. verum, Helianthemum vulgaris, Origanum vulgare, Phleum phleoides, nummularium, Hypochaeris maculata, Planta- Plantago lanceolata, Primula veris, Rhytidi- 75 Julia Stigenberg et al. Ent. Tidskr. 138 (2017) elevation 36 m. Sun exposed ridge side facing south east. Origanum vulgare, Ranunculus bul- bosus, Festuca ovina, Helictotrichon pratense, Helianthemum nummularium, Vincetoxicum hirundinaria, Juniperus communis, Geranium sanguineum, Filipendula vulgaris, Galium ver- um, Plantago lanceolata, Fragaria viridis. Collected material was sorted by participants jointly, and then identification responsibilities were divided among us as follows: AL & HV: Symphyta, MF: , Mymaridae, Mymarommatidae, Aphelinidae, Encyrtidae, Torymidae, , Trichogrammatidae, Figitidae, Dryinidae, , MF & JB: Platygastroidea, Ceraphronoidea, Tetracampi- dae, JB: , , Pteromali- dae, MF & HV: Cynipidae, : , AR: Ctenopelmatinae, NJ & AR: Ophioninae, Pim- plinae, , , IÖ: Cryptinae, , NJ: , Ichneumo- ninae, , Anomaloninae, Chrysidi- dae, , Pompilidae, , Crabroni- Figure 4. Niclas Eklund assisting with putting up Malaise dae, AL: Formicidae. Identification effort varied trap nr 4. In this trap we collected two of the new species to substantially between groups. Sweden; Gnamptodon decoris (Förster, 1862) and Entedo- Hymenoptera material is mainly deposited nomphale bulgarica Boyadzhiev & Triapitsyn, 2007. at the Swedish Museum of Natural History. In- Niclas Eklund hjälper till med uppsättningen av Malais- cidentally collected material of other taxa was efälla nr 4. I denna fälla fångades två nya arter för Sverige; either passed on to specialists or deposited at Gnamptodon decoris (Förster, 1862) och Entedonomphale the Campus Gotland division of Uppsala Uni- bulgarica Boyadzhiev & Triapitsyn, 2007. versity.

Other included collecting efforts In order to extend our snapshot of the island’s adelphus triquetrus, Satureja vulgaris, Sesleria Hymenoptera fauna, we decided to include a uliginosa, Thalictrum sp., Veronica chamae- small amount of other recently collected materi- drys, Vincetoxicum hirundinaria; the meadow als. Ole Lønnve had visited the island in May northwards rising towards the drier crest. 2008, collecting Symphyta, mainly by sweep- Malaise trap #5, 57.28369°N 17.97001°E, el- ing. In 2011, a group of entomologists visited evation 34 m. Just where the edge of the ridge the island in the tradition of the 80s visits and turned into northeast-facing slope. Patches of made a snapshot inventory, but this time includ- juniper, St. Lucie cherry (Prunus mahaleb) and ing Malaise traps (Elmqvist 2012). From these Cotoneaster sp., otherwise dominated by herbs 2011 traps stems the first Karlsö material includ- and grass (Dactylis glomerata, Festuca rubra, ing a broad range of parasitic Hymenoptera. The Filipendula vulgaris, Fragaria viridis, Galium following year, Niklas Johansson made an in- verum, Helianthemum nummularium, Helicto- ventory of the aculeate wasps, during which a trichon pratense, Phleum sp., Plantago lanceo- lot of larger parasitic wasps were collected but lata, Polygala vulgaris, Scabiosa columbaria, not included in the published report (Johansson Thalictrum sp., Vincetoxicum hirundinaria). 2013) and for the most part had not been identi- Malaise trap #6, 57.284763°N 17.970253°E, fied until now.

76 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö Results and Discussion Table 1. Number of genera, species and new species for The primary focus of the inventory was the par- the country found in 37 families of Hymenoptera during the Stora Karlsö snapshot inventory and the collecting events asitic wasps. Two thirds of the parasitic made in recent years. Note that many specimens could not families recorded in Sweden were represented be identified to species-level and thus we divided in two in the inventory. We recorded twelve species of separate columns for genera and species identified. The parasitic wasps that are new to Sweden, and sev- actual number of genera and species are higher. eral other species that have rarely been collected Antalet arter och nya arter för Sverige funna i 37 stekel- (Appendix 1). At least 15 subfamilies each of familjer under Stora Karlsöinventeringen 2014 samt i de the families Braconidae and inventeringar som gjorts åren precis före. En del stekar are represented in the material and about 200 kunde inte identifieras till art, därav två separata kolumner species of parasitoid wasps and 14 sawflies are för släkten och arter. Det faktiska antalet släkten och arter reported for Stora Karlsö for the first time. A är högre. rather sizeable amount of chalcid wasps (Chal- No of genera No of species No of species cidoidea) and figitid wasps (Figitidae) were col- Family identified identified new for Sweden lected, among these some rare species. While for 1 1 0 13 17 1 small-bodied wasps, the Malaise traps provided Gasteruptiidae 1 2 1 the major part of the catch, it became obvious 8 8 1 that middle-sized and larger parasitic wasps Proctotrupidae 4 4 0 were very patchily distributed in the landscape. 12 2 0 Whereas the major part of the heath was dry Ceraphronidae 2 0 0 Megaspilidae 3 0 0 and sweeping here was for the most part unre- Mymarommatidae 1 1 0 warding, spots that retained moisture were far Mymaridae 6 0 0 more productive. Especially the bog (Myren) Aphelinidae 3 0 0 on the south eastern part of the island was a Encyrtidae 3 0 0 Torymidae 2 0 0 particularly productive locality, as most of the Eupelmidae 2 4 0 ichneumonids in the inventory were collected Chalcididae 1 1 0 there, in particular . A large number Eulophidae 20 31 1 of those came from the immediate vicinities of 7 5 0 a large Salix tree. Included in the identifications Tetracampidae 1 1 0 Trichogrammatidae 1 0 0 are also various catches, obtained mainly with Cynipidae 4 4 0 Malaise traps by MF in 2011, and a rich assem- Figitidae 9 20 0 bly of ichneumonids and gasteruptiids feeding Braconidae 32 23 2 on flowering Laserpitium near Hien, collected Ichneumonidae 62 87 6 Dryinidae 1 1 0 by NJ in 2012. Below we present information Bethylidae 1 3 0 on new and interesting findings. Chrysididae 1 3 0 Formicidae 9 22 0 Gasteruptiidae Vespidae 6 11 0 No gasteruptiids were collected in 2014, but the Pompilidae 9 13 0 10 17 0 material from 2012 contained a few specimens, 1 1 0 including one which represents a species previ- Halictidae 3 10 0 ously not reported from Sweden: Gasteruption Andrenidae 1 6 0 opacum Tournier, 1877 (Fig. 5a). According to 2 6 0 6 10 0 Fauna Europaea this species is recorded from Apidae 2 4 0 Norway, but the record is based on an error (Frode Ødegaard, pers. comm.) and the clos- Total 250 318 12 est true known occurrences are from Germany and Poland. It is a bee parasitoid, but nothing is known on the particular habits of this species.

77 Julia Stigenberg et al. Ent. Tidskr. 138 (2017) a

Figure 5. Some rare and interest- ing wasps that were found on Stora Karlsö: – a) Gasteruption opacum, – b) Omphale sp., – c) Spilomicrus rufit- arsis, – d) Entedonomphale bulgarica. Några rara och intressanta steklar funna på Stora Karlsö: – a) Gasterup- tion opacum, – b) Omphale sp., – c) Spilomicrus rufitarsis, – d) Entedo- nomphale bulgarica.

Proctotrupoidea from the Nordic countries until recently, when Malaise traps and sweeping produced a rich it was discovered in Skåne, Södermanland and material of Diapriidae and Proctotrupidae, but a Uppland as well as in Norway and Finland large part of it is identified to -level only. within a relatively short time span (Forshage & The most remarkable record was Spilomicrus Karlsson 2015; Hansen 1997; Vikberg & Vark- rufitarsis (Kieffer, 1911) of Diapriinae, new to onyi 2006; Fredrik Ronquist & Roy Danielsson, Sweden (Fig. 5c). It is recorded from the British pers. comm.). Through the Swedish Malaise Isles but not from northern Europe. According Trap Project (SP) it has now been found all over to Nixon (1980) it is a rare species. Sweden, sometimes in large numbers, but it is very interesting to note that it occurs even on Platygastroidea & Ceraphronoidea relatively isolated localities like Stora Karlsö. Platygastroidea and Ceraphronoidea have only This indicates long-distance dispersal capaci- been identified to genus-level at this point, ex- ties, similar to other tiny parasitoids (Antolin & cept for two characteristic scelionines: Xeno- Strong 1987). merus ergenna Walker, 1836 that has charac- Its biology is still unknown. Interpolation teristic bottlebrush antennae, and the extremely from the biology of its family suggests that it flattenedTelenomus danubialis Szelenyi, 1939. is most likely an egg parasitoid, and Psocoptera have been suggested as a possible hostgroup. Chalcidoidea: Aphelinidae, Encyrtidae, Mymari- dae, Pteromalidae, Torymidae, Trichogrammatidae Tetracampidae These families have only been identified to ge- Epiclerus nomocerus (Masi, 1934) was the only nus-level or less at this point. representative of this rarely collected family. It is one of the least frequently collected tetracam- Mymarommatidae pids in Sweden, as it has so far not been found The spectacular and tiny Mymaromma anom- in the SP (MF, unpublished). The Swedish dis- alum (Blood & Kryger, 1922) was not known tribution seems to be restricted to Öland and 78 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö Gotland. Wasps of this genus are parasitoids of isander need to be widened and its intraspecific leaf-mining flies (Agromyzidae) but nothing is variation quantified to encompass the above- known about the biology of this particular spe- mentioned characters. cies. Eupelmidae Eulophidae In addition to several specimens of two quite A single male of Entedonomphale bulgarica common species with brachypterous females Boyadzhiev & Triapitsyn, 2007 (Fig. 5d) was (Eupelmus vesicularis and Merostenus excava- identified from Malaise trap #4. This recently tus), we also caught two macropterous speci- described species is a new record for Sweden; mens (one female and one male) of Eupelmus it was hitherto known only from Bulgaria (Boy- annulatus Nees, 1834 which was unknown from adzhiev & Triapitsyn, 2007) but its habitus is Sweden until very recently (Gibson & Fusu very distinctive. Its host is unknown, but other 2016). members of this genus are larval parasitoids of a single family of thrips (Thysanoptera: Phlaeo- Cynipoidea thripidae). Notable cynipoids include an undescribed spe- Another remarkable eulophid, a single male cies of Trybliographa (previously known from specimen of Omphale (Fig. 5b), was swept on other places in Sweden, not uncommon), the alvar vegetation of Stordal. It is quite similar relatively rare species Sarothrus brevicornis to Omphale isander (Walker, 1839) by having Thomson, 1877 and Callaspidia defonscolom- an asymmetrically shaped scape, remarkably bei (Dahlbom, 1842). These species are all para- long marginal setae of the forewing, and dark sitoids on fly larvae, the two former possibly of brown body colouration. However, the present Anthomyiidae (biology unknown in both cases specimen differs from O. isander, as described but this is a guess based on closely related spe- by Hansson & Shevtsova (2012), by having the cies), and the latter of aphid-predacious Syrphi- scape widest above the middle, the pedicel ap- dae and Chamaemyiidae. proximately as long as the maximal width of the scape, a pale antenna and a pale clypeus which Braconidae contrasts with the surrounding dark frons, and The impression regarding braconids on Stora the absence of a medial tooth on the clypeus. Karlsö is that the species richness is rather mod- The pale colour of the clypeus distinguishes it est. We managed to collect representatives of from most other species in this genus, and due 17 out the 29 subfamilies that occur in Sweden to this character, the present specimen runs be- (JS pers. comm.) The largest groups Alysiinae, tween O. clypealis (Thomson, 1878) and O. Aphidiinae, Microgastrinae and Braconinae are parma Hansson & Shevtsova, 2012 in the key abundant in specimens but less diverse regard- to European Omphale given by Hansson & ing species composition. This scarcity notwith- Shevtsova (2012). However, the specimen dif- standing, two new species for Sweden were fers from O. clypealis by having the clypeus not found: Bracon rozneri Papp, 1998 (Braconi- yellowish white but instead yellowish brown nae) and Gnamptodon decoris (Förster, 1862) (concolourous with the mandibles), and the (Gnamptodontinae). Gnamptodon decoris (Fig. forewing with only 6 admarginal setae; it dif- 6a) was caught in Malaise trap #4. It occurs in fers from O. parma by a closed speculum and most European countries, the closest to Sweden a slender stigmal vein which is less than half as are Germany, UK, Finland and Russia. It is not wide apically than basally. From both species, noted for Denmark or Norway. Gnamptodon- it differs by the scape that is widest above the tinae contains some of the smallest Braconidae, middle and abruptly narrowed in its apical third, usually hardly longer than 1 mm. They are ex- by the long marginal setae of the forewing, and clusively parasitoids of leafmining caterpillars the dark brown body colouration. It is possible of the Nepticulidae (Lepidoptera) on herbs and that this is just an aberrant O. isander male; this low shrubs. They are recognized by the peculiar would mean that the species boundaries of O. basal elevation of the second metasomal tergite 79 Julia Stigenberg et al. Ent. Tidskr. 138 (2017) a b

c d

Figure 6. Some exquisite wasps that were found on Stora Karlsö during this snapshot inventory: – a) Gnamptodon decoris, – b) Lissonota picticoxis, – c) Heterischnus filifor- mis, – d) Virgichneumon dumeticola. Några utsökta steklar funna på Stora Karlsö: – a) Gnampto- don decoris, – b) Lissonota picticoxis, – c) Heterischnus filiformis, – d) Virgichneumon dumeticola.

(van Achterberg 1983). Three Bracon species lected, Chelonus pusillus Szépligeti, 1908, was (identified by Konstantin Samartsev) were col- recently found as new to Sweden in the SP (iden- lected, of which 38 specimens of B. pectoralis tified by Aurel Lozan 2007 but yet unpublished). were swept close to a Salix shrub. These Bracon The few Opiinae specimens collected all be- species are probably not that uncommon though long to the genus Opiostomus, which has not rarely identified. Another notable species is yet been reported from Sweden but is far from the rare Meteorus sibyllae Stigenberg, 2011 uncommon, and will be presented in a coming (Euphorinae). It is not new to Sweden but new paper by Dave Karlsson. to Gotland, not so frequently collected and not common in Swedish collections. Ichneumonidae The only representative of col- A remarkably large part of the species of Ichneu- 80 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö monidae were collected exclusively by sweep- Plectochorus iwatensis (Uchida, 1928), this be- ing, rather than Malaise trapping, and most sig- ing the northwesternmost record of the species, nificantly from a single locality on the island, and Mesochorus tipularius Gravenhorst, 1829. the bog Myren. Especially the spider parasitoids Ophion brevicornis Morley, 1915 and Enicospi- in the Pimplinae were numerous there. As a lus inflexus (Ratzeburg, 1844) (Ophioninae) are contrast, the Banchinae was the only group that new to the list of Swedish species, but this is appeared to be common also on drier higher due to name confusion, which is currently be- grounds on the island. ing sorted out by Björn Cederberg (and NJ) in Habrocampulum biguttatum (Gravenhorst, Sweden (forthcoming), and by Broad & Shaw 1829) (Anomaloninae) is apparently a rare spe- in Britain (2016). cies, and we are not aware of any recent Swed- ish records of it. It attacks various pine-living Symphyta moths (Schmiedknecht 1908). Erigorgus latro The sawfly fauna of the island had not previ- (Schrank, 1781) (Anomaloninae) is a vari- ously been thoroughly explored. Coulianos able species with several varieties described (2010) found two gall-making species. We have (Schmiedknecht 1908). It is regarded as rare identified most of the relatively few specimens throughout its distribution range and the moth we collected, all of which belong to the family Diloba caeruleocephala is mentioned as a po- Tenthredinidae, but the time of the year of the tential host. Diloba caeruleocephala feeds on inventory was not optimal for sawfly collection. Prunus spp., which suggests an association with A more appropriate time for a comprehensive the enormous shrubberies of Prunus mahaleb inventory of sawflies would be from late spring that are commonly found around Suderslätt. to early summer, when the flight periods of Lissonota picticoxis Schmiedeknecht, 1900 most species take place. Sawfly larvae are her- (Banchinae) (Fig. 6b) is known from Finland bivorous and more or less specialized in their and is here reported for the first time from Swe- host-plant use, so that many sawfly species only den. No host records are known but the species attack a single or a few species of host plant. De- of this genus are usually caterpillar parasitoids. spite its isolation and relatively small area, be- The first specimen ofBathythrix maculata (Hel- cause of the extraordinary limestone-associated len, 1957) (Cryptinae) recorded from Sweden flora and warm climate, Stora Karlsö thus might was swept at the Myren bog. The distribution host more sawfly species not commonly seen in of this species ranges from Iran in the southeast Sweden than those that we have recorded below: to France in the west and Finland in the north. It One of the collected sawfly species, Athalia has been reared from chrysomelid beetle larvae cornubiae Benson, 1931 (Fig. 7) (Athaliinae (Sawoniewicz 1980). (often placed in ), is a new record for For Platylabus heteromallus (Ichneumoni- Sweden. The larva feeds on white stonecrop (Se- nae), our specimen represents the first certain dum album) a plant that was observed next to the record from Scandinavia, whereas Virgichneu- trap on the southern shore where the species was mon dumeticola (Fig. 6d) was recently reported collected. The species has a Holarctic distribu- as new to Sweden from the SP (Riedel & Mag- tion, but prior to this it has not been found in the nusson 2014). Heterischnus filiformis (Graven- Nordic countries (Taeger & Blank 2011). It does horst, 1829) (Fig. 6c) is a member of the Phaeo- not seem to be very common and is probably genini which previously has been recorded from restricted to one host plant (Gradwell 1957). most neighboring countries including Norway Several other species of Athalia were col- and Finland. Several females and males were lected. When conditions are favourable, the spe- swept at Myren. The species has also been col- cies of this genus potentially have more than one lected recently from the Swedish mainland (Nor- generation per year (Benson 1952), which may rköping, Östergötland Leg. Håkan Andersson, explain why they were numerous in August. This det. Niklas Johansson) and is here presented as is also the case for the curled rose sawflyAllan - new to Sweden. Two species of the subfamily tus cinctus, which is often seen as adults in the Mesochorinae are reported as new to Sweden, autumn (Hartig 1837). Of the subfamily Nema- 81 Julia Stigenberg et al. Ent. Tidskr. 138 (2017)

Figure 7. The sawfly Athalia cornubiae Benson, 1931. Also a new species to Swe- den, collected in Malaise trap nr 5. Photo by Artur Larsson. Växtstekeln Athalia cornubiae Benson, 1931 vars larver le- ver på vit fetknopp (Sedum album) var också en art för Sverige. Den fångades i Malaisefälla nr. 5

tinae, we found a couple of species of Pristiph- son 2013). Johansson (2013) noted that the pro- ora, which may also have 2 or 3 generations and portion of the parasitic aculeates was relatively can be found in the autumn (Chawner & Pea- low. Some aculeates new to the fauna of Stora cock 1923, Viitasaari 2002). Another possibil- Karlsö were also recorded. The solitary wasp ity of the high numbers in August might be that Stenodynerus bluethgeni Van der Vecht, 1971 they hibernate in the egg stage, complete their seems to be restricted to alvar habitats. Recent larval development during early summer and fly Swedish records are all from first-class dry habi- as imagines during late summer or spring. This tats on Öland. It is absent from the rest of the is rare among the sawflies, but is for example Nordic countries and its closest occurrences are known for the European pine sawflyNeodiprion in the Netherlands and central Germany (Abe- sertifer (Pschorn-Walcher, 1965). nius 2012). On Gotland, the most recent record As could be expected, a totally different set of was from Buttle 1972, until a female was found species were observed by Ole Lønnve when he at Suderslätt in 2012 (Johansson 2013) as the visited the island for a couple of days at the end first record in 40 years. But since then this spe- of May in 2008 (Appendix). The juniper-feeding cies has also been found in a few localities on Monoctenus obscuratus (Diprionidae) was then the Gotland main island. Two specimens of the observed in large numbers all across the island, sphecid wasp Rhopalum gracile Wesmael, 1848 whereas the other sawfly species were observed were swept at a reed bed at the bog Myren. This at the bog Myren and utilize host plants like Fili- small wasp primarily inhabits stems of reed and pendula ulmaria (Empria) or different species is found in reed-beds and lush marshes (Blösch of ferns (Strongylogaster macula), commonly 2000). Due to its habitat preferences, it is prob- seen at the edge of the bog. ably overlooked but the records are new to the province of Gotland (Hellqvist et al 2014). Lasi- oglossum zonulum Smith, 1848, is a medium The 2012 inventory of the island’s aculeate sized halictid bee which is quite common on the fauna showed, together with older records, adjacent areas on Gotland. One male was swept that Stora Karlsö harbour several rare aculeate on a dry meadow just west of Myren. The spe- wasps that should be protected and an ecosys- cies is new to the fauna of Stora Karlsö. tem with a unique species composition (Johans- Another interesting observation was that the 82 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö rare Gotlandic subspecies of Andrena marginata very effective and standardised way to sample Fabricius, 1777; A. marginata nigrescens Auriv- biodiversity, but to draw meaningful conclu- illius, 1903, which has previously been recorded sions in terms of island biogeography within from the island (Nilsson 2011), could not be one particular time window, a replicated study found although the timing of the inventory was may be required, setting up the same number of just right with respect to its flight period, in some replicated traps in a comparable vegetation on places along with lots of flowering Succisa pra- both islands as well as on Gotland and on the tensis, which is the species’ preferred host plant. mainland within the same time period. Coelioxys obtusispina Thomson, 1872 is a The fact that we found a number of new spe- cleptoparasite of the bee Megachile lagopoda cies to Sweden and a number of rare species (Linnaeus, 1761). This species has a very re- probably reflects the poor knowledge of the stricted world distribution and is currently hymenopteran fauna in general, and we deem known only from Gotland. Up until the 1950s it likely that any similar collection and identi- it was also present in some other parts of Swe- fication effort on any relatively rich spot in the den, in Östergötland and Uppland, but since country would produce new and rare species in these populations seem to have disappeared the a similar magnitude, as shown by the evidence species is currently to be regarded as a Gotland from the Swedish Malaise Trap Project (unpub- endemic. From Stora Karlsö, Olov Lundblad lished). Our snapshot of the Stora Karlsö wasp collected the species in the 1950s and it was re- community may not on its own illuminate ques- discovered in 2012 (Johansson 2013). tions concerning ecological patterns but even so, it is a gleaming glint of the fauna, filled with Conclusions notable surprises. Since both climatic conditions and vegetation Our study thus provides an easy recipe on may be very different from other nearby lo- how the knowledge of Swedish biodiversity calities, an island fauna can hold big surprises can be substantially augmented: By a short and and seem very peculiar. This certainly seems intensive collecting expedition, making use of to be the case with Stora Karlsö. Our material the synergistic effect of different experts. We necessarily contains only a small subsample of strongly recommend that other interesting lo- the island’s fauna, representing a limited time calities in the country are subjected to similar window. Numerous singletons in the material efforts by temporary teams of determined ento- suggest the observed species richness is still mologists with slightly different target groups. far from the factual. Thus, it would be prema- Provided that a permit can be acquired from the ture to draw any conclusions in terms of island authorities, there are still many hidden species biogeography. Further studies will be needed to discover; and from the point of view of na- to address community assembly topics such as ture conservation, we can only devise effective the proportions of specialists vs. generalists, protection measures if we know what is there to and comparison with mainland communities in be protected. similar habitat patches; or indications of locali- ty-specific adaptations of ecological strategies. Acknowledgements Any study focussing on proportions of special- The inventory was financed by Karlsö Jagt och Djur- ists versus generalists will, however, require skyddsförening AB. Håkan Elmqvist was helpful dur- some knowledge of the biology of each species, ing the preparations and Niclas Eklund is thanked for which is unfortunately non-existent for many assisting when putting up the Malaise traps. Andrew poorly-known groups of Hymenoptera. Liston is thanked for helping with identification of Symphyta, Konstantin Samartsev identified Braco- It may be further interesting to assess the ninae, Dave Karlsson identified Opiinae, Christer effect of discontinued sheep grazing on biodi- Hansson helped with identification of Eulophidae, versity by a faunal comparison between Stora and Reijo Jussila, Matthias Riedel, Erich Diller and Karlsö and its closest neighbour, Lilla Karlsö, Gavin Broad helped with Ichneumonidae identifica- where sheep grazing remained continuous up to tions. Bertil Widbom kindly loaned us stereo micro- the present day. Particularly Malaise traps are a scopes during the inventory 83 Julia Stigenberg et al. Ent. 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Long-distance dis- norvegica Series B 44: 81-82. persal by a parasitoid (Anagrus delicatus, Mymari- Hanski, I. 1999. Metapopulation Ecology. – Oxford dae) and its host. – Oecologia 73: 288-292. University Press. Arrhenius, O. 1921. Species and area. – Journal of Hanski, I. 2016. Habitat Connectivity, Habitat Continu- Ecology 9: 95-99. ity, and Metapopulations in Dynamic Landscapes. Benson, R.B.1952. Symphyta Section (b). – In: Hand- – Oikos, 87: 209-219. books for the identification of British Insects. Roy- Hansson, C. & Shevtsova, E. 2012. Revision of the Eu- al Entomological Society of London. London. UK. ropean species of Omphale Haliday (Hymenoptera, 137 pp. Chalcidoidea, Eulophidae). – ZooKeys 232: 1-157. Blösch, M. 2000. Die Grabwespen Deutschlands. Sphe- Hartig, . 1837. Die Aderflügler Deutschlands mit be- cidae s.str., Crabronidae. Lebensweise, Verhalten, sonderer Berücksichtigung ihres Larvenzustandes Verbreitung. – Die Tierwelt Deutschlands, 71 Teil. und ihres Wirkens in Wäldern und Gärten für En- Goecke & Evers, Keltern. tomologen, Wald- under Gartenbesitzer. Die Fami- Boyadzhiev, P.S. & Triapitsyn, S.. 2007. Description lien der Blattwespen und Holzwespen nebst einer of a new species of Entedonomphale (Hymenop- allgemenen Einleitung zur Naturgeschichte der tera: Eulophidae) from Bulgaria, with notes on E. Hymenopteren. – Haude und Spener, Berlin 1:i- carbonaria. – Revue Suisse de Zoologie 114(4): xiv, 1-416. 735-741. Hedgren, S. 2005. Restaurering av alvarmiljöer Broad, G. & Shaw, M. 2016. The British species of på Stora Karlsö. – Slutrapport för projekt nr. Enicospilus. – European Journal of Taxonomy 187: LIFE00NAT/S/7118. Länsstyrelsen Gotlands Län. 1-31. Hellqvist, S., Abenius J. & Norén, L. 2014. Provins- Broqvist, . & Magntorn, O. 1989. Inventering av träd- förteckning för de svenska arterna i familjerna Am- och buskvegetationen på Stora Karlsö. – Läns- pulicidae, Sphecidae och Crabronidae (Hymenop- styrelsen Gotlands Län, Naturvårdsenheten, Visby. tera). – Entomologisk Tidskrift 125: 77-94. 122 pp. Högmo, O. 1988. Myror på Stora Karlsö. – FaZett 1: Chawner, E.F. & Peacock, A.D. 1923. Observations 11-12. on the Life-Histories and Habits of Allantus pal- Johansson, N. 2013. Gaddsteklar på Stora Karlsö 2012. lipes, Spin., and Pristiphora pallipes, Lep. (Hym. – Rapporter om natur och miljö nr 2013: 2. Läns- Tenth.). – The Entomologist 56: 125-128. styrelsen Gotlands län. http://www.lansstyrelsen. Coulianos, C-C. 2010. Gallbildningar funna på Stora se/gotland/SiteCollectionDocuments/Sv/Pub- Karlsö. – Rindi 30: 3-17. likationer/Natur-och-milj%C3%B6/2013/2013- Elmqvist, H. 2012. Insektinventering på Stora Karlsö 02-gaddsteklar.pdf 10-14 augusti 2011. – Karlsöbladet 2012 (1): 5-6. Karlsson, D., Pape, T., Johanson K.A, Liljeblad, J. & Eriksson M.O.G. & Rosén E. 2008. Management of Ronquist, F. 2005. Svenska Malaisefälleprojektet, Natura 2000 habitats. 6280 *Nordic alvar and pre- eller hur många arter steklar, flugor och myggor cambrian calcareous flatrocks. – European - Com finns i Sverige? – Entomologisk Tidskrift 126: 43- mission 53. Forshage, M. & Karlsson, D. 2015. Taxonomic units Laforest, B.J., Winegardner, A.K., Zaheer, O.A., Jef- in the SP: Third-tier Chalcidoidea – http://www. fery, N.W., Boyle, E.E. & Adamowicz, S.J. 2013. stationlinne.se/sv/forskning/the-swedish-malaise- Insights into biodiversity sampling strategies trap-project-sp/taxonomic-units-in-the-sp/third- for freshwater microinvertebrate faunas through tier-chalcidoidea/ bioblitz campaigns and DNA barcoding. – BMC Fukami, T. 2015. Historical contingency in community ecology 13:13. assembly: integrating niches, species pools, and Larje, R. 2008. Stora Karlsö – a tiny Baltic island with a priority effects. – Annual Review of Ecology Evo- puzzling past. – Terra australis 29: 265-279. lution and Systematics 46: 1-23. Linnaeus, C. 1745. Öländska och Gothländska Resa riksens högloflige Ständers befallning förrättad

84 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö Åhr 1741. Med Anmärkningar uti Oeconomien, (07.12.2011). – Digital Entomological Information, Natural-Historien, Antiquiteter &c med åtskillige Müncheberg Figurer. – Stockholm och Upsala: hos Gottfried Vikberg, V. & Várkonyi, G. 2006. Paljepistiäinen (Pa- Kiesewetter. laeomymar anomalum) todettu Etelä-Suomesta Linnman, S. 1965. Om skalbaggsfaunan på Karlsöarna. (Hymenoptera: Mymarommatoidea: Mymarom- – Opuscula Entomologica 30: 183-210. matidae). – Sahlbergia 11: 42-48 MacArthur, R.H. & Wilson, O.E. 1967. The Theory of Viitasaari, M. 2002. The suborder Symphyta of the Island Biogeography. – Princeton University Press. Hymenoptera. – In: Viitasaari, M. (ed.) Sawflies I Malaise, R. 1937. A new insect trap. – Entomologisk Tremex Press. Helsinki. Finland. Tidskrift 58: 148-160. With, K.A. 2016. Are landscapes more than the sum of Nilsson, L.A. 2011. Andrena marginata nigrescens their patches? – Landscape Ecology 31: 969-980. Aurivillius (Sw. Gotländskt guldsandbi), a third subspecies of bee from Gotland (Hymenoptera, Apoidea, Andrenidae). [Andrena marginata nigres- Svensk sammanfattning cens Aurivillius Gotländskt guldsandbi, en tredje gotländsk bi-underart.] – Entomologisk Tidskrift Stekelfaunan på Stora Karlsö utanför Got- 132: 25-31. land är överlag dåligt känd trots att den gan- Nixon, G.E.J. 1980. Diapriidae (Diapriinae). – Hand- ska märkvärdiga gaddstekelfaunan undersökts books for the Identification of British Insects vol tidigare. För att få mer kunskap om denna fauna VIII, part 3(di). Royal Entomological Society of och se hur mycket man kan lära sig med en kort London, 55 pp. gemensam insats genomförde vi, åtta personer, Pschorn-Walcher, H. 1965. The ecology of Neodiprion sertifer Geoffr. (Hym.,Diprionidae) and a review en inventering under fem dagar i augusti 2014 of its parasite complex in Europe. – Technical Bul- med främst genom handplockning och håvning letin of the Commonwealth Institute of Biological i vegetation. Vi hade också Malaisefällor Control 5: 33-97. uppsatta under 22 dagar. Främst inriktade vi Riedel, M. & Magnusson, P. 2014. A contribution to the oss mot parasitsteklar och växtsteklar. Vi fick Swedish fauna of (Hymenoptera, ihop ca 200 arter parasitsteklar och 14 växt- Ichneumonidae), with descriptions of the female steklar som rapporteras för första gången från of Cratichneumon armillatops Rasnitsyn 1981 and the male of Aoplus biannulatorius (Thunberg Stora Karlsö. Fynd från ett par andra besök på 1824). – Entomofauna 35: 433-448. ön har också inkluderats i resultaten här – av Roslin, T., Várkonyi, G., Koponen, M., Vikberg, V. Ole Lønnve 2008, Mattias Forshage m fl 2011, & Nieminen, M. 2014. Species-area relationships Niklas Johansson 2012 och Carl-Cedric Cou- across four trophic levels – decreasing island size lianos vid flera tillfällen. Totalt 12 arter (inklu- truncates food chains. – Ecography 37: 443-453. derar ett fynd av NJ 2012) arter rapporteras som Sawoniewicz, J. 1980. Revision of European species nya för Sverige: bladstekeln Athalia cornubiae of the genus Bathythrix Foerster (Hymenoptera, Ichneumonidae). – Annales Zoologici 35: 319-365. Benson, 1931, bistekeln Gasteruption opacum Schmiedeknecht, O. 1908. Anomaloninae. – In: Opus- (Tournier, 1877), hyllhornstekeln Spilomicrus cula Ichneumonologica, Fasc. XIX, p 1457-1520. rufitarsis (Kieffer, 1911), finglansstekeln Ent- Blankenburg i Thür. edonomphale bulgarica Boyadzhiev & Triapit- Schwenke, . 1999. Revision der europäischen Meso- syn, 2007, bracksteklarna Bracon roznari Papp, chorinae (Hymenoptera, , Ichneu- 1998 och Gnamptodon decoris (Förster, 1862), monidae). – Spixiana Suppl. 26: 1-124. och följande brokparasitsteklar: Bathythrix Shaw, M.R. & Hochberg, M.E. 2001. The neglect of parasitic Hymenoptera in insect conservation strat- maculata (Hellén, 1957), Heterischnus filifor- egies: The British fauna as a prime example. – Jour- mis (Gravenhorst, 1829), Lissonota picticoxis nal of Insect Conservation 5: 265-263. Schmiedeknecht, 1900, Mesochorus tipularius Stolpe, H. 1882. Förteckning öfver svenska myror, pre- Gravenhorst, 1829, Ophion brevicornis Mor- liminärt meddelande. – Entomologisk Tidskrift 2: ley, 1915, och Plectochorus iwatensis (Uchida, 127-151. 1928). Detta visar hur kunskapsläget för svensk Struwe, I. 1989. Inventeringsvecka på Stora Karlsö – om biologisk mångfald kan ökas massivt under en entomologisk glädje. – Graphosoma 4(1): 14-16. Taeger, A. & Blank, S.M. 2011. ECatSym - Elec- kort men intensiv inventering. Vi rekommen- tronic World Catalog of Symphyta (Insecta, Hy- derar starkt att fler platser i Sverige inventeras menoptera). Program version 3.9, data version 38 på liknande vis.

85 Julia Stigenberg et al. Ent. Tidskr. 138 (2017) ------ - x ------x x - x x x - - x ------x - - x - - x - x - - x x - - x x - - x - x x all - 3 - - - 2 3 x x 3 4 3 x 5 2 x 1 2 x 4 - 5 5 4 x - x x > > 4 all 4 3 x x x x

(Haliday, 1839) (Haliday, (Thomson, 1859) (Linnaeus, 1758) (Haliday, 1839) (Haliday, (Möller, 1882) (Möller, 1911) (Kieffer, Szelenyi, 1939 1911) (Kieffer, (Latreille, 1805) Walker, 1836 Walker, (Thomson, 1859) (Thomson, 1859) (Curtis, 1831) Thomson, 1859 spp. sp. spp. spp. sp. spp. spp. sp. sp. 1 sp. 2 sp. 3 sp. 4 sp. 5 sp. Appendix, continued. DIAPRIIDAE Belytinae Belyta depressa Cinetus sp. Aclista Pantolyta sp. Diapriinae Psilus fuscipennis * Spilomicrus rufitarsis Basalys Basalys Basalys Basalys Basalys Trichopria sp. Trichopria Trichopria oogaster Trichopria Trichopria aequata Trichopria Trichopria wasmanni Trichopria Trichopria cf. tenuicornis Trichopria verticillata Trichopria PROCTOTRUPIDAE Proctotrupes bistriatus Proctotrupes gravidator Codrus spp. Pschornia spp. Phaneroserphus calcar Phaenoserphus cf. viator PLATYGASTRIDAE Inostemma Leptacis Platygaster Amblyaspis Synopeas sp. Scelioninae s. lat. Gryon sp. Trimorus spp. Trimorus Trissolcus Trissolcus Scelio sp. Telenomus Telenomus Telenomus danubialis Telenomus Anteris sp. Xenomerus ergenna CERAPHRONIDAE Aphanogmus Ceraphron spp. MEGASPILIDAE Megaspilus

Other records Other ------Coulianos 1989 Coulianos - - - - - Coulianos 2009 Coulianos - - - - -

(2008) Lønnve

-

(2011) Forshage

- - -

Johansson (2012) Johansson - - -

sweep net & handp. & net sweep - x - - x - x x - - x - - x - x - - - x - x - x - x - x - x

no trap Malaise -

4 - - - 5 5 - - - - 2 1,4,5 x ------

Viitasaari & Vikberg, 1985 & Vikberg, Viitasaari (Hartig, 1837) (Linnaeus, 1758) (Fallén, 1808) (Klug, 1917) (Gimmerthal, 1847) (Tournier, 1877) (Tournier, (Müller, 1776) (Müller, (Gmelin, 1790) (Geoffroy in Fourcroy, 1785) in Fourcroy, (Geoffroy (Fabricius, 1781) (Audinet-Serville, 1823) Benson, 1931 Linnaeus, 1758 (Cameron, 1883) (Klug, 1816) (Klug, 1815) Audinet-Serville, 1823 (Konow, 1896) (Konow, (Linnaeus, 1758) DIPRIONIDAE Monoctenus obscuratus Appendix. Hymenoptera taxa found on Stora Karlsö. Columns indicate where re - handp.” are from the pres - cords are obtained; “Malaise trap no” and “sweep net & ent inventory with trap no’s as in Fig. 1, ”x” = presence in unspecified trap(-s), “>” =presence in several traps.; references indicate records from earlier sources. New species for Sweden are marked with “*”. förstade två när fynden gjorts: anger Kolumnerna Karlsö. på Stora som hittas Steklar “>” fälla/-or ospec i förekomst = ”x” 1, Fig. i som fällnr. = siffra inventering: denna från = förekomt i fler fällor; kolumner med referenser anger litteraturkällor för fynd. Nya arter för Sveroge markeras med “*”. TAXON TENTHREDINIDAE Allantinae Apethymus serotinus Empria pallimacula Empria pumila Athaliinae Athalia cordata Athalia rosae * Athalia cornubiae Athalia circularis Blennocampa phyllocolpa Claremontia waldheimii Heterarthriinae aethiops pumilus Cladius pectinicornis Pristiphora pallidiventris bridgmanii Selandriinae Nesoselandria morio Strongylogaster macula * Gasteruption opacum Tenthredininae cf. atra Tenthredo GASTERUPTIIDAE

86 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö ------x ------x x - - - x - x x x x x - - x - - x - - x x x x x x x - - x - x x x x x - x - - x 2 5 x 1 - - - > 4 > 5 4 4 2 - 1 1 - - - - 5 5 x x 1 2 - - 1 - - x

(Zetterstedt, 1838) Boucek, 1958 (Walker, 1839) (Walker, (Walker, 1834) (Walker, (Nees von Esenbeck, 1834) (Haliday, 1844) (Haliday, (Walker, 1839) (Walker, (Nees von Esenbeck, 1834) (Dalman, 1818) (Nees, 1834) (Nees von Esenbeck, 1834) (Boyer de Fonscolombe, 1832) (Walker, 1839) (Walker, (Spinola, 1808) (Linnaeus, 1758) Walker, 1838 Walker, (Walker, 1839) (Walker, (Zetterstedt, 1838) Boucek, 1958 (Walker, 1838) (Walker, (Walker, 1839) (Walker, Walker, 1838 Walker, (Walker, 1839) (Walker, (Ashmead, 1904) (Walker, 1838) (Walker, Spinola, 1811 (Walker, 1839) (Walker, sp. sp. 1 sp. 2 sp. 3 sp. 4 sp. 5 sp. sp. Appendix, continued. Entiinae Euderus albitarsis Elachertus isadas Elachertus lateralis Cirrospilus lyncus Cirrospilus vittatus Diglyphus begini Diglyphus isaea Elasmus flabellatus Elasmus sp Dicladoderus euryalus Eulophus larvarum Hemiptarsenus ornatus Hemiptarsenus unguicellus Necremnus metalarus Pnigalio soemius Sympiesis gordius Sympiesis gregori Sympiesis sereicornis cf. orithyia Aprostocetus Aprostocetus Aprostocetus Aprostocetus Aprostocetus Crataepus marbis Tamarixia Tamarixia Tamarixia pubescens Tamarixia monesus Tamarixia upis Tamarixia Tetrastichus atratulus Tetrastichus PTEROMALIDAE indet. Pteromalidae Asaphinae Asaphes sp. Asaphes suspensus Miscogastrinae Halticoptera patellana Trichomalus Trichomalus campestris Trichomalus Cyrtogaster sp. Trichomalus cf. lonchaeae Trichomalus Callitula bicolor Pachyneuron

Other records Other ------

(2008) Lønnve

------

Forshage (2011) Forshage x - x - x x -

Johansson (2012) Johansson ------

sweep net & handp. & net sweep x x - - - - x - x ------x x - - - x ------x x x - - x x - x x x

no trap Malaise x

all 3 3 4 4 4 > > > 1 - x 6 2 2 all 4 x 1 x 5 - - - 4 2 - - 4 - - -

(Erdös, 1954) (Westwood, 1833) (Westwood, (Walker, 1839) (Walker, (Blood & Kryger, 1922) (Blood & Kryger, (Zetterstedt, 1838) Nees von Esenbeck, 1834 (Dalman, 1820) (Retzius, 1783) (Walker, 1839) (Walker, Dalman, 1820 Hansson & Shevtsova, 2012 (Olivier, 1791) (Olivier, (Walker, 1839) (Walker, sp. sp. spp. spp. sp. sp. spp. Conostigmus Appendix conuuied TAXON Dendrocerus spp. MYMAROMMATIDAE Mymaromma anomalum MYMARIDAE Alaptus sp. Anagrus spp. Anaphes sp. Erythmelus Gonatocerus Ooctonus APHELINIDAE Aphelinus Centrodora sp. Coccophagus Encyrtidae indet. ENCYRTIDAE Encyrtidae Copidosoma sp. Bothriothorax sp. ?Aphycoides sp. ?Aphycoides TORYMIDAE spp. Torymus Megastigmus EUPELMIDAE Eupelmus vesicularis Eupelmus cf. annulatus Eupelmus urozonus Eupelmus sp. ( males ) Merostenus excavatus CHALCIDIDAE Haltichella rufipes EULOPHIDAE Entedontinae Neochrysocharis formosa Neochrysocharis chlorogaster Chrysocharis nephereus Chrysocharis pubicornis Chrysocharis polyzo Boyadzhiev & Triaptsyn, 2007 *Triaptsyn, Entedonomphale bulgarica Boyadzhiev & Ionympha carne Omphale sp, aetius-group isander Omphale nr. Omphale incognita

87 Julia Stigenberg et al. Ent. Tidskr. 138 (2017) ------x - x - - - x - x - x - - x ------x - x - x x x x x x - x - - - x - x x x x x - x - x - x - - - x - x - - - - x x x - x x - x - - x x - x x x x 1 - > > - - 4 x 5 x 3 x > 4 - x x x - x all

(Nees von Esenbeck, 1834) (Nees von Esenbeck, 1812) Curtis, 1833 Fischer, 1960 Fischer, (Förster, 1862) (Förster, (Nees von Esenbeck, 1834) Haliday, 1833 Haliday, (Wesmael, 1835) (Wesmael, (Nees von Esenbeck, 1834) Reinhard, 1862 Nees, 1834 (Nees von Esenbeck, 1834) (Nees von Esenbeck, 1834) Szépligeti, 1908 (Linnaeus, 1758) Stigenberg, 2011 Wesmael, 1838 Wesmael, Ruthe, 1861 (Wollaston, 1858) (Wollaston, sp. spp. Appendix, continued. Alysiinae Chorebus sp. Aspilota spp. Alysia sp. Orthostigma Dapsilarthra sp. Phaenocarpa sp. Dacnusa spp. Dinotrema spp. Aphidiinae Aphidius sp. Diaeretus sp. Pauesia sp. Praon sp. Blacinae Blacus armatulus Braconinae * Bracon rozneri Papp, 1998 Bracon exhilarator Bracon pectoralis Cheloninae Chelonus pusillus Adelius subfasciatus Doryctinae Spathius exarator Heterospilus separator Ontsiria antica Euphorinae Meteorus ruficeps Meteorus ictericus Meteorus cf. abdominator Meteorus sibyllae Leiophron pallidistigma Townesilitus bicolor Townesilitus Gnamptodontinae * Gnamptodon decoris Helconinae Diospilus morosus Hormiinae Hormius moniliatus Macrocentrinae Macrocentrus infirmus Macrocentrus thoracicus Macrocentrus Microgastrinae indet. Microgastrinae Microgastrinae Apanteles sp.

Other records Other - - - Coulianos 1989, 2009 1989, Coulianos Coulianos 1988, 2009 Coulianos 1989, ------

(2008) Lønnve

------

(2011) Forshage

x -

Johansson (2012) Johansson ------

sweep net & handp. & net sweep x - - x - x x x x x x ------x x x - - x - x - - - x x - x - -

no trap Malaise 1,5 x - - -

- 2 5 - - 1 4 x 4 > 2 > - 5 5 4 - 5 1 x - - - 5 5

(Linnaeus, 1758) Kieffer, 1908 Kieffer, (Dahlbom, 1842) (Hartig, 1841) (Hartig, 1841) (Hartig, 1841) (Kieffer, 1904) (Kieffer, Westwood, 1833 Westwood, (Dalman, 1818) (Thomson, 1862) (Hartig, 1840) Thomson, 1862 Thomson, 1877 (Masi, 1934) (Cameron, 1879) Cameron, 1889 (Kieffer, 1901) (Kieffer, (Thomson, 1862) (Giraud, 1860) (Walker, 1835) (Walker, (Curtis, 1838) ) 1833) ( MF 2011 (Westwood, (Giraud, 1860) (Kieffer, 1902) (Kieffer, (Linnaeus, 1758) sp. sp. 1 sp. 1 ( males ) sp. 1 TETRACAMPIDAE Epiclerus nomocerus Appendix conuuied TAXON TRICHOGRAMMATIDAE Trichogramma CYNIPIDAE Aulacidea pilosellae Diplolepis eglanteriae Diplolepis rosae Neuroterus quercusbaccarum FIGITIDAE Anacharitinae Anacharis immunis Anacharis eucharioides Xyalaspis armata Aspicerinae Callaspidia defonscolombei Charipinae Alloxysta cf. macrophadna Alloxysta cf. pleuralis Alloxysta fulviceps Alloxysta cf. citripes Alloxysta arcuata Alloxysta cf. tscheki Alloxysta Alloxysta cf. victrix Eucoilinae Kleidotoma cf. psiloides Kleidotoma cf. myrmecophila Kleidotoma Kleidotoma cf. affinis Trybliographa cf. diaphana Trybliographa Trybliographa agaricola Trybliographa Trybliographa cf. trichopsila Trybliographa Trybliographa Trybliographa Didyctium cf. nigriclavum Figitinae Figites cf. validicornis Sarothrus brevicornis BRACONIDAE Bassus sp.

88 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö ------x ------x x - x x x x x x - - x x - x x x x - x x x x x x x x x x x x x x x x x x x x x x x x x - x - 1 - - - x 2 5 2 5 1 - 2 - - - - x ------1 - - 1 -

(Förster, 1850) (Förster,

(Linnaeus, 1758) (Gmelin, 1790) (Gravenhorst, 1829) (Thomson, 1884) (Thomson, 1884) (Thunberg, 1824) (Panzer, 1809) (Panzer, (Schrank, 1781) (Uchida, 1928) (Berthomieu, 1910) (Gravenhorst, 1829) Wesmael, 1845 Wesmael, Gravenhorst, 1829 (Gravenhorst, 1829) (Gmelin, 1790) (Schrank, 1802) Hartig, 1838 Holmgren, 1860 Linnaeus, 1758 Holmgren, 1858 (Wesmael, 1845) (Wesmael, (Gravenhorst, 1829) (Thomson, 1890) (Fabricius, 1775) (Gravenhorst, 1829) Tischbein, 1873 Tischbein, Teunissen, 1943 Teunissen, (Thunberg, 1824) Wesmael, 1853 Wesmael, (Fabricius, 1771) Curtis, 1833 (Thunberg, 1824) (Thunberg, 1822) (Gravenhorst, 1829) (Gravenhorst, 1829) (Gravenhorst, 1829) (Gravenhorst, 1829) (Gravenhorst, 1829) Wesmael, 1845 Wesmael, (Fabricius, 1775) Megacara vagans Appendix, continued. Phygadeuontina sp. Orthizema cf. hadrocerum Zoophthorus nr. plumbeus Zoophthorus nr. Diplazontinae Diplazon laetatorius Diplazon tetragonus Diplazon scutatorius Diplazon sp. Promethes sulcator Homotropus signatus Homotropus dimidiatus Tymmophorus erythrozonus Tymmophorus Syrphoctonus tarsatorius Syrphophilus tricinctorius Enizemum ornatum Enizemum nigricorne Ichneumoninae indet. Ichneumoninae Ichneumoninae Ichneumon sarcitorius Ichneumon suspiciosus Ichneumon simulans Cratichneumon flavifrons Chasmias motatorius Homotherus locutor Barichneumon peregrinator Virgichneumon dumeticola Virgichneumon Platylabus rufus Platylabus pallidens Platylabus curtorius Platylabus heteromallus Platylabus orbitalis Platylabus dolorosus * Heterischnus filiformis Centeterus rubiginosus Phaeogenes melanogonos Dirophanes fulvitarsis Dicaelotus pumilus Diadromus subtilicornis Aethecerus sp. Mesochorinae Mesochorus gemellus Mesochorus olerum Mesochorus rubeculus Mesochorus semirufus * Mesochorus tipularius * Plectochorus iwatensis

Other records Other ------

(2008) Lønnve

- -

(2011) Forshage

-

(2012) Johansson

------

sweep net & handp. & net sweep - - - x x - x x - - x x x x x x x x x x x - x - x - x x x x x x x - x x

no trap Malaise x

2 - - 1 - - x 1 - 1 1 - - - - - 2 - x x - x ------x

(Gravenhorst, 1829) (Nees von Esenbeck, 1834) (Thunberg, 1824) Haliday, 1839 Haliday, (Fabricius, 1793) (Gravenhorst, 1829) Gravenhorst, 1829 (Wesmael, 1838) (Wesmael, (Hellén, 1957) Gravenhorst, 1829 Schmiedeknecht, 1900 (Müller, 1776) (Müller, (Poda, 1761) (Gmelin, 1790) Gravenhorst, 1829 Gravenhorst, 1829 (Schrank, 1781) (Gravenhorst, 1829) (Spinola, 1808) Thomson, 1889 Thomson, 1889 (Panzer, 1804) (Panzer, spp. sp. sp. sp. Appendix conuuied TAXON Opiinae Opiostomus Orgilinae Orgilus Rhyssalinae Oncophanes minutus Rogadinae Aleiodes bicolor Aleiodes circumscriptus ICHNEUMONIDAE Anomaloninae Habrocampulum biguttatum Erigorgus cf. latro Banchinae indet. Banchinae Banchinae Apophua sp. Exetastes adpressorius Exetastes gracilicornis Glypta caudata Glypta dentifera Glypta bifoveolata Glypta fronticornis Lissonota Lissonota coracina * Lissonota picticoxis Lissonota cf. accusator Campopleginae indet. Campopleginae Campopleginae Ctenopelmatinae indet. Ctenopelmatinae Ctenopelmatinae Rhinotorus sp. Rhinotorus Cryptinae indet. Cryptinae Cryptinae Aritranis sp. Cryptus dianae s. lat. Ischnus inquisitorius Stenarella domator Atractodes croceicornis Bathythrix fragilis * Bathythrix maculata Dichrogaster aestivalis Gelis Gelis areator

89 Julia Stigenberg et al. Ent. Tidskr. 138 (2017) - Norén - Norén Norén - Frisk 1972 Frisk Högmo Högmo Norén 2009, Högmo 2009, Norén Högmo - 1988 Högmo, Hagman 2013 Hagman - - Frisk 1971, Högmo 1971, Frisk Frisk 1972 Frisk Högmo1988 - Högmo Norén, Frisk 1972 Högmo Frisk 1988, Norén, Bartsch Högmo Frisk 1972, Högmo 1972, Frisk - Högmo Frisk 1972, Högmo - - - - Norén - Norén Norén Norén Norén Norén Lundblad Lundblad Lundblad Lundblad Lundblad ------x x ------x - - - - x - x - x - x - x - x - x - x - x x - - - x - x - - x - x - x - - - x - x - x - x - x - x - x - x - x - x - x - x - x - x - x - x - - x ------> - x - x ------x - all - - 4 ------x ------

)

Ødegaard & Soon, 2015

(Fabricius, 1793)

(Müller, 1776 (Müller, van der Vecht, 1971 van der Vecht, (Fabricius, 1793) (Linnaeus, 1761) (Wesmael, 1836) (Wesmael, (Linnaeus, 1758) (Fabricius, 1793) (Latreille, 1802) (Latreille, 1802) (Nylander, 1846) (Nylander, (Linnaeus, 1758) Linnaeus, 1758 (Curtis, 1829) (Spinola, 1808) ( Fabricius, 1775 Seifert, 1992 (Wesmael, 1851) (Wesmael, (Dahlbom, 1843) Linsenmaier, 1959 Linsenmaier, (Schiødte, 1837) (Jurine, 1807) Latreille, 1798 Bondroit, 1920 (Fabricius, 1793) Förster, 1850 Förster, Nylander, 1846 Nylander, Seifert, 1992 (Haupt, 1927) Latreille, 1798 Meinert, 1861 Emery, 1895 Emery, (Linnaeus, 1758) Elmes, 1978 (Thomson, 1861) Finzi, 1926 (Linnaeus, 1758) (Förster, 1850) (Förster, Linnaeus, 1758 Shuckard, 1836 Forel, 1886 (Fabricius, 1782)

Appendix, continued. Bethylus fuscicornis Bethylus boops FORMICIDAE Myrmica schenki CHRYSIDIDAE subcoriacea Chrysis borealis Paukkunen, Myrmica sabuleti Myrmica rubra Myrmica ruginodis Myrmica hirsuta Myrmica lonae Myrmica cf. vandeli Myrmecina graminicola Solenopsis fugax Leptothorax muscorum Leptothorax acervorum Tetramorium caespitum Tetramorium Lasius alienus Lasius paralienus Lasius psammophilus Lasius flavus Formica fusca Formica cunicularia Formica polyctena Formica clara Camponotus ligniperda Temnothorax tuberum Temnothorax VESPIDAE germanica Vespula vulgaris Vespula Dolichovespula saxonica Stenodynerus bluethgeni Eodynerus quadrifasciatus Symmorphus murarius scoticus Ancistrocerus parietinus Ancistrocerus trifasciatus POMPILIDAE rufa Arachnospila spissa cinctellus

Other records Other ------

(2008) Lønnve

- - -

(2011) Forshage

- - x -

Johansson (2012) Johansson ------

sweep net & handp. & net sweep - x x x - x x x x x - x x x x x x x x x x x - - x - - x x x x x x - - x - x x -

no trap Malaise

3 - x 2 - x ------2 - - x - 5 x - - x - 6 - 3 - x x )

(Hartig, 1838))

(Perkins, 1957) (Haliday, 1838) (Haliday, ( planatus (Thomson, 1877) Retzius, 1783 Ruthe, 1855 (Thunberg, 1822) (Fonscolombe, 1854) (Dalman, 1818) (Linnaeus, 1758) (Panzer, 1800) (Panzer, (Holmgren, 1860) (Müller, 1776) (Müller, Morley, 1915 Morley, (Förster, 1860) (Förster, (Ratzeburg, 1844) (Holmgren, 1860) (Gravenhorst, 1829) (Holmgren, 1860) (Gravenhorst, 1829) (Gravenhorst, 1829) (Gravenhorst, 1829) Jussila, 1965 (Gravenhorst, 1829) Holmgren, 1856 Thomson, 1877 (Schrank, 1802) ( annulatus Kiss (Gravenhorst, 1829) Gravenhorst, 1829 Appendix conuuied TAXON Stictopisthus unicinctor Ophioninae * Ophion brevicornis Pimplinae Pimpla spuria Pimpla flavicoxis Pimpla hypochondriaca Pimpla contemplator Itoplectis aterrima Itoplectis alternans Apechthis quadridentata Apechthis compunctor Endromopoda detrita Scambus eucosmidarum Scambus calobatus

Scambus inanis Scambus vesicarius Zaglyptus varipes Schizopyga circulator Clistopyga rufator Sinarachna pallipes Tromatobia ornata Tromatobia Orthocentrinae indet Orthocentrinae Orthocentrinae Stilbopinae Stilbops sp Tersilochinae Phradis morionellus Tryphoninae indet. Tryphoninae Tryphoninae Phytodietus variegatus Netelia sp Netelia melanura Ctenochira haemosterna Exenterus ictericus Grypocentrus cinctellus Thymaris tener DRYINIDAE Aphelopus melaleucus Dryinidae indet. Dryinidae BETHYLIDAE Bethylus cephalotes

90 Ent. Tidskr. 138 (2017) Hymenoptera fauna on Stora Karlsö Norén - Norén Norén Norén Norén Norén Norén Norén Norén - Norén Lundblad Norén Norén - - - Lundblad, Norén Lundblad, Lundblad - - Norén - Lundblad, Norén ------x - x - x - - - - x - x - x - - x - x - x - x - x - x - x - x x - - - x - x - x - x - x - x ------

(Panzer, 1806) (Panzer, (Kirby, 1802) (Kirby, Thomson, 1872 (LePeletier, 1841) (LePeletier, Perez, 1902 (Kirby, 1802) (Kirby, (Kirby, 1802) (Kirby, (Linnaeus, 1761) Nylander, 1852 Nylander, Fabricius, 1777 (Panzer, 1798) (Panzer, (Thomson, 1872) Nylander, 1852 Nylander, LePeletier, 1841 LePeletier, Smith, 1842 (Kirby, 1802) (Kirby, Nylander, 1852 Nylander, (Linnaeus, 1758) (Linnaeus, 1761) (Panzer, 1799) (Panzer, Fabricius, 1781 Eversmann, 1852 Nylander, 1852 Nylander, Appendix, continued. ANDRENIDAE Andrena haemorrhoa (Fabricius, 1781) Andrena labiata Andrena carantonica Andrena fucata Smith, 1847 Andrena nigroaenea Andrena marginata COLLETIDAE Hylaeus hyalinatus Hylaeus communis Hylaeus brevicornis Hylaeus confusus Hylaeus annularis Colletes floralis MEGACHILIDAE Megachile lagopoda Megachile versicolor Smith, 1844 Megachile willlugbhiella Hoplosmia spinulosa APIDAE Anthophora furcata Anthophora quadrimaculata Bombus lucorum Coelioxys obtusispina Coelioxys elongata Osmia aurulenta rapunculi Hoplitis mitis Hoplitis claviventris Bombus terrestris

Other records Other Lundblad Lundblad, Norén Lundblad, Lundblad Lundblad Lundblad Lundblad Lundblad - - - - Norén - - Norén Norén Norén - Norén Norén Norén - - Norén - - Norén Norén Norén Norén - - Norén Lundblad - Norén

(2008) Lønnve

- - - -

Forshage (2011) Forshage ------

Johansson (2012) Johansson ------

sweep net & handp. & net sweep - x - - x - - - x - x - x x - - x - x - x - x - x - x x - x - - x - x - x - x - x - x x - x - x x - - x - - x - x

no trap Malaise - -

------2 - - - - > - - - - - 2 ------

(Morawitz, 1866) (van der Linden, 1829) (Kirby, 1802) (Kirby, (Spinola, 1808) (Fabricius, 1804) (Fabricius, 1781) (Kirby, 1802) (Kirby, (Smith, 1848) Smith, 1851 (Scopoli, 1763) (Schuckard, 1837) Say, 1824 Say, (Fabricius, 1781) (Linnaeus, 1767) Blüthgen, 1931 (Linnaeus, 1758) (Fabricius, 1793) (Fabricius, 1804) (Thomson, 1870) (Fabricius, 1775) (Fabricius, 1793) (Scopoli, 1763) Thomson, 1870 Beaumont, 1945 (Dahlbom, 1843) (Linnaeus, 1758) (Linnaeus, 1758) (Gmelin, 1790) (LePeletier, 1832) (LePeletier, (Dahlbom, 1843) Smith, 1853 Beaumont, 1945 Wesmael, 1852 Wesmael, Dahlbom, 1844 (Linnaeus, 1758) (Fabricius, 1787) rufipes nigerrimus Appendix conuuied TAXON Anoplius concinnus fasciatellus crassicornis hyalinata maculata variegatus CRABRONIDAE Nysson maculosus Gorytes laticinctus Gorytes quadrifasciatus continuus Ectemnius cavifrons dimidiatus Crossocerus elongatulus Crossocerus distinguendus Rhopalum coarctatum Rhopalum gracile Trypoxylon medium Trypoxylon attenuatum Trypoxylon Trypoxylon minus Trypoxylon niger inornata Psenulus concolor Mimesa lutaria SPHECIDAE Ammophila sabulosa HALICTIDAE Lasioglossum morio Lasioglossum albipes Lasioglossum leucopus Lasioglossum villosulum Lasioglossum zonulum Halictus confusus Halictus tumulorum Halictus eurygnathus Sphecodes crassus Sphecodes ephippius

91