An illustrated key to the cuckoo wasps (Hymenoptera, Chrysididae) of the Nordic and Baltic countries, with description of a new species

Juho Paukkunen; Alexander Berg; Villu Soon; Frode Ødegaard; Paolo Rosa

doi:10.3897/zookeys.548.6164

Review Article

An illustrated key to the cuckoo wasps (Hymenoptera, Chrysididae) of the Nordic and Baltic countries, with description of a new species

Juho Paukkunen^‡, Alexander Berg^§, Villu Soon^|, Frode Ødegaard^¶, Paolo Rosa^#

‡ Finnish Museum of Natural History, Helsinki, Finland

§ Lund University, Lund, Sweden

| University of Tartu, Tartu, Estonia

¶ Norwegian Institute for Nature Research - NINA, Trondheim, Norway

# Unaffiliated, Bernareggio, Italy

Corresponding author: Juho Paukkunen ( juho.paukkunen@helsinki.fi )

Academic editor: Michael S. Engel

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Paukkunen J, Berg A, Soon V, Ødegaard F, Rosa P (2015) An illustrated key to the cuckoo wasps (Hymenoptera, Chrysididae) of the Nordic and Baltic countries, with description of a new species. ZooKeys 548: 1-116. https://doi.org/10.3897/zookeys.548.6164

ZooBank: urn:lsid:zoobank.org:pub:D5D7B51E-5AC6-460D-9B3C-7584E46F9B3F

Abstract

The Chrysididae are a group of cleptoparasitic and parasitoid aculeate wasps with a large number of rare and endangered species. The taxonomy of this group has long been confusing due to the similarity of species and extensive intraspecific variation. We present for the first time a comprehensive dichotomous key for all 74 species found in the Nordic and Baltic countries. In addition to diagnostic characters, information on the distribution and biology of each species is also presented. A new species, Chrysis borealis Paukkunen, Ødegaard & Soon, sp. n. is described on the basis of specimens collected from Fennoscandia. Chrysis gracillima Förster, 1853 is recorded as new to the Nordic and Baltic countries.

Keywords

Morphology, distribution, phenology, host species, Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden, Chrysis borealis sp. n.

Introduction

Chrysidid wasps, also known as cuckoo wasps, represent one of the largest families of aculeate Hymenoptera within the superfamily Chrysidoidea. More than 2.500 species are known worldwide (Aguiar et al. 2013) and approximately 490 of these have been recorded from Europe (Mitroiu et al. 2015). The species richness decreases towards the north of Europe, and a total of 74 species have been found in Fennoscandia, Denmark and the Baltic countries (Paukkunen et al. 2014). Cuckoo wasps, excluding Amiseginae and Loboscelidiinae, which are not present in Europe, are well known for their bright metallic colours and cleptoparasitic or parasitoid lifestyle. In northern Europe, species of the subfamily Chrysidinae parasitise solitary wasps (Vespidae and Crabronidae) and solitary bees (Megachilidae), whereas species of Cleptinae attack tenthredinid and diprionid sawflies. Despite their attractive appearance, chrysidids have a reputation for being a taxonomically difficult group, and the biology of several species is still poorly known.

A detailed history of cuckoo wasp research in the Nordic and Baltic countries was presented recently by Paukkunen et al. (2014). More than 200 publications including information on chrysidids have been published from the region, but most of them consist of only scattered records in poorly circulated journals or reports. Only a few authors have conducted more extensive faunistic studies at a national or wider scale. Faunistic surveys, including identification keys, have been compiled by Dahlbom (1829, 1831), Thomson (1862, 1870) and Aurivillius (1911) in Sweden, by Borries (1891) in Denmark, and by Sahlberg (1910) and Hellén (1920) in Finland. The monographs of Dahlbom (1845, 1854) also include keys and descriptions of several Nordic species. A simple key to the Swedish genera was later presented by Landin (1971). The faunistic studies from Estonia (Soon 2004), Latvia (Tumšs and Maršakovs 1970) and Lithuania (Orlovskytė et al. 2010) did not include identification keys of chrysidids. In recent decades, species determination of chrysidids in the Nordic and Baltic countries has mainly relied on the works of Linsenmaier (1951, 1959, 1997), Morgan (1984), Kunz (1994) and van der Smissen (2010), which focus primarily on the central European fauna, but include most of the North European species.

Scattered notes on the biology of European cuckoo wasps have been published by several authors in numerous articles and reports, and these data have been compiled by e.g. Kunz (1994) and Rosa (2006). An important contribution to the knowledge of hosts of North European species was recently made by Pärn et al. (2014) in Estonia. The Nordic open access databases of entomological observations, Artportalen (http://www.artportalen.se) in Sweden, Artskart (http://artskart.artsdatabanken.no) in Norway and Hyönteistietokanta (http://hyonteiset.luomus.fi) in Finland provide extensive sources of information on the phenology and habitats of cuckoo wasps. Many unpublished records of hosts, habitats and phenology can also be found in public and private cuckoo wasp collections.

As most publications with information on the identification and biology of the Nordic and Baltic cuckoo wasps are scattered, outdated and/or difficult to find or use, there is a need for a new comprehensive key for the North European species including biological information. Cuckoo wasps include an exceptionally large number of red-listed and endangered species in the Nordic countries, which also highlights the importance of their reliable identification (Cederberg et al. 2010, Hansen et al. 2010, Paukkunen 2010).

The aim of this study is to present a simple dichotomous identification key for the Nordic and Baltic species, and to compile all relevant and reliable information on their distribution, abundance and biology, including phenology and host species from publications and collections. The key will hopefully arouse more interest in chrysidids among entomologists, and provide a basis for further, more detailed studies on the distribution, biology and morphology of North European species.

Material and methods

The geographic area covered by the study includes the Nordic and Baltic countries, which are located in northern Europe (Fig. 1). The nomenclature and arrangement of the taxa follows Paukkunen et al. (2014) and Rosa et al. (2015), and the morphological terminology is based on Morgan (1984) and Kimsey and Bohart (1991), with a few exceptions. Most notably, mesosoma is used instead of thorax, metasoma instead of abdomen or gaster, mesoscutum instead of scutum and mesoscutellum instead of scutellum. These exceptions are made in order to harmonise modern general Hymenoptera terminology (Hymenoptera Anatomy Consortium 2015). The following abbreviations are used: T = tergite, S = sternite and F = flagellomere (= flagellar segment, pseudosegment of flagellum). Numbers are used for antennal and metasomal segments, for example, F2 refers to the second flagellomere. The key is mainly based on the works of Linsenmaier (1997) and van der Smissen (2010). In order to keep the key simple and concise, only easily visible and relatively constant characters have been selected. Several new diagnostic characters have also been found and included in the key. In addition to figures of morphological details, one dorsal habitus picture of an entire specimen is presented for each genus.

The species treatments consist of the following information: name, synonymy, diagnosis, distribution and biology. Only the more common synonyms and erroneously interpreted names, which have been used in connection with cited records from the study area, are presented below the valid name of the taxon. If the currently used name differs from the original combination, it is added to the synonymic list with a citation of the study, in which the rearrangement was made. The abundance of each species is estimated using the scale 1) very common (more than 5000 records), 2) common (ca 1000–5000 records), 3) relatively common (ca 500–1000 records), 4) relatively rare (ca 200–500 records), 5) rare (ca 10–200 records), 6) very rare (less than 10 records). This estimation is mainly based on collected material and therefore it essentially shows how commonly a species is collected, but might not accurately indicate its actual abundance in nature. A summary of the distribution of chrysidid species in the Nordic and Baltic countries is presented in Table 3.

The biology section includes information on the habitat, flight season and host species. The presented information on the distribution, abundance and biology has been compiled from published literature, entomological databases and several public and private collections, as well as our own observations. Host species, plants and habitats that are not found in the Nordic and Baltic countries are usually not mentioned. The most important studied collections are listed below:

LMSZ Museum of Zoology, University of Latvia; Riga, Latvia

MZH Finnish Museum of Natural History, University of Helsinki; Helsinki, Finland

MZLU Zoological Museum, Lund University; Lund, Sweden

NHRS Swedish Museum of Natural History; Stockholm, Sweden

NMLS Natur-Museum Luzern; Luzern, Switzerland

NUM NTNU University Museum, Trondheim, Norway

TUZ Natural History Museum, University of Tartu; Tartu, Estonia

ZMAA Zoological Museum, Åbo Akademi University; Turku, Finland

ZMUC Zoological Museum, University of Copenhagen; Copenhagen, Denmark

ZMUO Zoological Museum, University of Oulu; Oulu, Finland

ZMUT Zoological Museum, University of Turku; Turku, Finland

Lists of examined material have not been included in the species treatments due to the large number of studied specimens. Accurate data is given only if a species is recorded for the first time from a country. Some information about the examined material has been published earlier by Paukkunen et al. (2014), and most of the Finnish data is openly accessible through the Finnish Biodiversity Info Facility (http://laji.fi). Data on DNA barcoded specimens are available at the Barcode of Life Data System (http://www.boldsystems.org, Ratnasingham and Hebert 2007).

Morphological measurements were prepared using an ocular micrometer on a Wild M5 and a Leica MZ75 stereomicroscope. All pictures were prepared by Alexander Berg, if not otherwise specified. The photos were taken with a Canon6D camera, using a Schneider-Kreuznach Componon-S 50 mm f2.8 and Schneider-Kreuznach Componon 28 mm f1.4 enlarger lenses extended on Pentacon M42 bellows. A Proxxon KT-70 microstage was used for photo stepping and Zerene Stacker v1.04 for stacking the photos.

In order to use the key successfully, specimens should be properly mounted or pinned with both the dorsal and ventral surfaces of the metasoma visible. In the Chrysis ignita and C. fasciata species-groups, the mandibles of both sexes should be opened, genital capsules of males should be extracted and ovipositors of females everted. Colouration of specimens collected with traps containing liquid preservatives, softened using hot water or having been kept in sunlight for a long time, can deviate from the original colouration. Additionally, the colour of fresh and liquid preserved specimens can change when they are dried, most notably greenish shades turn bluish in dry specimens. Geographical variation in colouration is also observed in many species, whereby northern specimens tend to be darker than southern ones.

Distinguishing the sexes of chrysidids can be difficult if the telescope-like ovipositor of the female is not exserted, or the genital capsule of the male has not been extracted. In males, the third metasomal sternite is completely flat and the semitransparent membranous posterior margin of the fourth sternite is usually visible. In females, the third sternite is generally thicker posteriorly and the posterior margin of the fourth sternite is opaque. Additionally, a slender needle-like structure (formed by the first valvulae) can be seen on the tip of the ovipositor in females. This structure is visible even if the ovipositor is not fully exerted.

Figure 1.

Map of the study area.

Figures 2–10.

Tarsal claw: 2 Pseudomalus triangulifer 3 Holopyga generosa 4 Hedychrum nobile 5 Hedychridium roseum 6 Chrysis longula. Forewing: 7 Holopyga generosa 8 Hedychridium ardens 9 Pseudospinolia neglecta 10 Chrysis impressa, a and c radial sector vein, b medial vein, d medial cell.

Systematics

Key to chrysidid genera of the Nordic and Baltic countries

1	Metasoma with four (female) or five (male) external tergites, ventral surface convex, colour anteriorly non-metallic red, posteriorly black, often with blue-green metallic reflections (Figs 11, 16, 17). Pronotum campanulate (Figs 11–13)	Cleptes Latreille (Cleptinae)
–	Metasoma with three (or four in Parnopes male) external tergites, ventral surface flat or concave, colour variable, often completely metallic, if non-metallic red then never posteriorly extensively black. Pronotum not campanulate. (Chrysidinae)	2
2	Metasoma with three (female) or four (male) external tergites (Fig. 209). T1 mostly metallic and subsequent tergites usually non-metallic red (Fig. 209). Posterior margin of metasoma with numerous small irregular teeth. Mouthparts (galea and glossa) strongly developed, longer than rest of head. Tegula large and broad, covering base of forewing and hindwing (Fig. 209)	Parnopes grandior (Pallas) (Parnopini)
–	Metasoma with three external tergites in both sexes. All tergites usually with metallic colour, and if non-metallic, then T1 also without metallic reflections. Posterior margin of metasoma without small irregular teeth. Mouthparts short. Tegula small, covering only base of forewing	3
3	Tarsal claw with one or more subapical teeth (Figs 2–5). Radial sector vein of forewing basally curved (Figs 7a, 8a). Head without preoccipital carina. T3 without subapical pit row and apical teeth. (Note: Elampus bidens (Förster), which might be found in North Europe in the future, has two apical teeth on T3, but also a tongue-like metascutellar projection, as in Fig. 38.) (Elampini)	4
–	Tarsal claw simple, without subapical teeth (Fig. 6). Radial sector vein of forewing basally straight (Figs 9a, 10a). Head with preoccipital carina, ending in hook (Figs 138–149). T3 with subapical pit row, with or without apical teeth. (Chrysidini)	10
4	Tarsal claw with single subapical tooth (Figs 4, 5). Temple rounded in dorsal view (Fig. 74). Medial cell of forewing with short setae	5
–	Tarsal claw with more than one subapical tooth (Figs 2, 3). Temple angular in dorsal view (Fig. 48). Medial cell of forewing with or without setae	6
5	Subapical tooth of tarsal claw almost as large as apex, so that tip appears forked (Fig. 4). T3 usually laterally with two small angular projections (Figs 61–63)	Hedychrum Latreille
–	Tarsal claw with very small submedial tooth, remote from apex (Fig. 5). T3 without angular projections	Hedychridium Abeille de Perrin
6	Medial cell of forewing with setae. Medial vein of forewing strongly curved (Fig. 7b). T3 without apical notch	Holopyga Dahlbom
–	Medial cell of forewing without setae. Medial vein of forewing only slightly curved (Fig. 8b). T3 usually with apical notch (Figs 21, 23, 25, 31, 32, 36, 44–46)	7
7	Metascutellum dorsally with large tongue-like projection (Fig. 38). Gena of female with row of dense short setae (Fig. 41)	Elampus Spinola
–	Metascutellum without projection. Gena without row of short setae	8
8	Mesoscutum with large punctures concentrated postero-medially between notauli (Fig. 27). Ventral margin of mesopleuron strongly projecting (Fig. 28). Head and mesosoma usually with long pubescence, setae twice as long as diameter of mid-ocellus	Pseudomalus Ashmead
–	Mesoscutum without punctures, with irregularly scattered punctures (Figs 20, 22, 24) or with punctures clumped along notauli (Fig. 35). Ventral margin of mesopleuron weakly projecting (Fig. 19). Head and mesosoma with short pubescence, setae not more than twice as long as diameter of mid-ocellus	9
9	Apical notch of T3 with thickened margin (Fig. 36). Lateral margin of T3 with concave depression prior to apical notch (Fig. 36). Mesoscutum with coarse punctation (Fig. 35). Metascutellum sharply elevated	Philoctetes truncatus (Dahlbom)
–	Apical notch of T3 without thickened margin (Figs 21, 23, 25). Lateral margin of T3 rounded or relatively straight, without concave depression prior to apical notch (Figs 21, 23, 25). Mesoscutum without or with finer punctation (Figs 20, 22, 24). Metascutellum not sharply elevated	Omalus Panzer
10	Radial sector vein of forewing ending before wing margin, at a distance approximately equal to the length of the pterostigma (Fig. 9c). Posterior margin of T3 without apical teeth	11
–	Radial sector vein of forewing extending to wing margin, or nearly so (Fig. 10c). Posterior margin of T3 with or without apical teeth	12
11.	Body entirely blue (Fig. 76). Medial cell of forewing without setae. Scapal basin with fine punctation	Spinolia unicolor (Dahlbom)
–	Body bicoloured, head and mesosoma blue-green, metasoma red (Fig. 75). Medial cell of forewing with setae. Scapal basin medially with fine cross-ridging	Pseudospinolia neglecta (Shuckard)
12	Posterior margin of T3 without apical teeth or angular prominences (Figs 206–208). Frons flat, without transverse frontal carina (Figs 201, 202). Male usually with F2 to F5 ventrally bulging (Fig. 204)	Chrysura Dahlbom
–	Posterior margin of T3 with apical teeth or angular prominences (Figs 78–88, 198), or rarely bluntly triangular or rounded without teeth (Chrysis gracillima, C. succincta and C. leachii) (Fig. 77). Frons with deep scapal basin and transverse frontal carina (Figs 150–154)	13
13	Posterior margin of T3 with three apical teeth, lateral teeth may be angular projections (Fig. 198). Black spots of S2 small and joined together into one central spot of variable shape (Fig. 199). Body entirely blue-green, sometimes blackish (Fig. 197)	Trichrysis cyanea (Linnaeus)
–	Posterior margin of T3 with different number of teeth (Figs 78–88), or rarely without teeth (Fig. 77). Black spots on S2 not as above. Body colour variable	Chrysis Linnaeus

I. Subfamily Cleptinae

This subfamily represents the most basally arising lineage of Chrysididae (Kimsey and Bohart 1991, Carpenter 1999). It is characterised by the following features: frons without scapal basin, pronotum narrowed submedially and campanulate in dorsal view (Figs 11–13), propodeum rectangular in profile, with horizontal dorsal surface, metasoma with four external segments in the female and five segments in the male (Figs 11, 16, 17), and metasomal venter convex. Cleptinae are parasitoids of sawfly prepupae (Hymenoptera: Symphyta) of the families Tenthredinidae and Diprionidae (Kimsey and Bohart 1991). The subfamily includes three genera, Cleptes Latreille, 1802, Cleptidea Mocsáry, 1904 and Lustrinia Kurian, 1955, of which only Cleptes is known from Europe (Kimsey and Bohart 1991, Móczár 1996). Currently, a total of 121 Cleptinae species are recognised worldwide (Wei et al. 2013, Arens 2014).

Genus Cleptes Latreille, 1802

Figs 11, 12–17

Cleptes Latreille, 1802: 316.

Note

Cleptes females search for tenthredinid and diprionid sawfly cocoons either on the host’s foodplant or on the ground beneath and lay one egg per cocoon (Morgan 1984). The emerging larva develops as an ectoparasitoid of the sawfly prepupa within the cocoon (Darling and Smith 1985, Kimsey and Bohart 1991). The genus consists of around 100 known species, the majority of which occur in the Holarctic Region (Kimsey and Bohart 1991, Wei et al. 2013, Arens 2014). A total of 27 species are known from Europe (Rosa and Soon 2012, Arens 2014) and three from the Nordic and Baltic countries (Paukkunen et al. 2014). We have divided the genus into species-groups according to Móczár (1997, 2001).

Figure 11.

Cleptes semiauratus ♀. Scale 1 mm. (Photo: Pekka Malinen).

Key to Cleptes species of the Nordic and Baltic countries

1	Pronotum posteriorly with furrow of foveae (Figs 11, 12). Female mesoscutum metallic golden-red (Fig. 11)	C. semiauratus (Linnaeus)
–	Pronotum without furrow of foveae (Fig. 13). Female mesoscutum black, sometimes with greenish reflections	2
2	Vertex with deep postocellar foveae (Fig. 14). Female head black without metallic reflections. Mesoscutum with regular punctation. Meso- and metatibiae yellowish. T2 with dense punctation (Fig. 16). T3 and T4 without metallic sheen, or only laterally with weak violet or bluish reflections (Fig. 16)	C. nitidulus (Fabricius)
–	Vertex without postocellar foveae (Fig. 15). Female head usually with metallic violet reflections. Mesoscutum with only a few punctures. Meso- and metatibiae brownish. T2 with sparse punctation (Fig. 17). T3 and T4 with strong blue or violet metallic sheen (Fig. 17)	C. semicyaneus Tournier

Cleptes nitidulus group

Cleptes nitidulus (Fabricius, 1793)

Figs 13, 14, 16

Ichneumon nitidulus Fabricius, 1793: 184.

Cleptes nitidula: Fabricius 1804: 154.

Diagnosis

Length 5–7 mm. Both sexes differ from C. semiauratus by not having a foveate furrow posteriorly on the pronotum (Fig. 13). The female also differs from C. semiauratus by its black head and mesoscutum, non-metallic yellow or orange pronotum, mostly metallic blue mesoscutellum, metanotum and propodeum, and non-metallic black apex of the metasoma. As opposed to the female, the head and mesosoma of the male are entirely metallic green and the apex of the metasoma has faint metallic reflections laterally (Fig. 16). Both sexes differ from C. semicyaneus by having pale brown or yellow (not dark brown) legs, denser punctation on the tergites (Fig. 16) and deep postocellar foveae on the vertex (Fig. 14).

Distribution

Denmark, Estonia, Finland, Latvia, Sweden. Rare. – West Palearctic: Europe and Turkey (Linsenmaier 1959), records from China are erroneous (Rosa et al. 2014).

Biology

Habitat: sparsely vegetated sandy areas, such as dry meadows and dunes (Morgan 1984). Occasionally found on flowers of Apiaceae (Heinrich 1964). Flight period: June to August. Host: Caliroa cerasi (Linnaeus) and Euura ribesii (Scopoli) (Tenthredinidae) (Dahlbom 1854, Morgan 1984).

Cleptes semicyaneus Tournier, 1879

Figs 15, 17

Cleptes semicyanea Tournier, 1879: 88.

Diagnosis

Length 4–7 mm. Both sexes resemble C. nitidulus superficially, but the legs are darker brown, the punctation of the tergites is sparser (Fig. 17) and the vertex does not have postocellar foveae (Fig. 15). The female differs also from C. nitidulus by having blue-violet metallic sheen posteriorly on the metasoma. In the male, this blue-violet sheen is more extensive (Fig. 17) than in C. nitidulus.

Distribution

Denmark, Norway, Sweden. Very rare. – Trans-Palearctic: from western Europe to Siberia (Irkutsk) (Móczár 1997).

Biology

Habitat: sparsely vegetated coastal sandy areas (Hansen et al. 2010, Fritz and Larsson 2010). Flight period: June to August. Host: unknown, possibly a sawfly living on creeping willows (Salix repens Linnaeus) (Ødegaard et al. 2009).

Figures 12–17.

Pronotum, dorsal view: 12 Cleptes semiauratus ♂ 13 C. nitidulus ♂. Head, dorsal view (arrow indicating postocellar fovea): 14 C. nitidulus ♂ 15 C. semicyaneus ♂. Metasoma, dorsal view: 16 C. nitidulus ♂ 17 C. semicyaneus ♂. Scale 1 mm.

Cleptes semiauratus group

Cleptes semiauratus (Linnaeus, 1761)

Figs 11, 12

Sphex semiauratus Linnaeus, 1761: 413.

Cleptes semi-auratus: Latreille 1802: 316

Cleptes pallipes Lepeletier, 1806: 119. Synonymised with C. semiauratus by Dahlbom (1854) and synonymy reinstated by Rosa et al. (2015).

Diagnosis

Length 5–8 mm. Both sexes differ from C. nitidulus and C. semicyaneus by having a transverse foveate furrow posteriorly on the pronotum (Figs 11, 12). The female differs also by its metallic golden red head, pronotum, mesoscutum and mesoscutellum (Fig. 11), and distinctly banded wings (Fig. 11). The head and mesosoma of the male are entirely metallic blue-green. The metasoma is anteriorly non-metallic red and posteriorly black with blue reflections in both sexes (Fig. 11).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – Trans-Palearctic/Holarctic? The general distribution is poorly known due to confusion of C. semiauratus with C. striatipleuris Rosa, Forshage, Paukkunen & Soon, 2015 (= C. semiauratus sensu Lepeletier, 1806) by several authors (Rosa et al. 2015). According to Móczár (2001), C. semiauratus has been found in the Palearctic, Nearctic and Oriental Regions (Sumatra). In the Nearctic and Oriental Regions the species has probably been accidentally introduced (Kimsey and Bohart 1991).

Biology

Habitat: forest margins and clearings, gardens and parks. Flight period: June to August. Host: Endelomyia aethiops (Gmelin), Euura ribesii (Scopoli) and Pristiphora incisa (Lindqvist) (Tenthredinidae) (Alfken 1915, Burger and Sobczyk 2011, V. Vikberg, pers. obs.). Several other tenthredinid sawfly species reported as hosts for C. semiauratus might actually represent hosts of C. striatipleuris.

II. Subfamily Chrysidinae

The majority of all chrysidids, about 80%, belong to this subfamily (Kimsey and Bohart 1991). Its members are characterised by a bright metallic colouration (with a few rare exceptions), three (or less commonly two or four) external tergites, and a concave or flat metasomal venter (with one rare exception). They are mainly nest parasites of solitary wasps and bees, although one genus, Praestochrysis, attacks prepupal moth larvae. The subfamily is distributed in all zoogeographical regions and consists of four tribes: Allocoelini (not present in Europe), Elampini, Chrysidini and Parnopini (Kimsey and Bohart 1991, Carpenter 1999).

Tribe Elampini

Chrysidid wasps of this tribe are characterised by three external metasomal tergites, the absence of a pit row or sublateral foveae on T3, and the usually dentate tarsal claw. The tribe has a worldwide distribution, though most of the genera and species occur in arid areas of the Holarctic Region. A total of 21 genera are recognised, seven of which are found in North Europe.

Genus Omalus Panzer, 1801

Figs 18, 19–25

Omalus Panzer, 1801: 13.

Note

Many authors have treated this genus in the broad sense and divide it into several subgenera (see summary by Rosa 2006). We follow the classification of Kimsey and Bohart (1991), whereby Elampus, Philoctetes and Pseudomalus are recognised as valid genera. Omalus sensu stricto is characterised by the following morphological features: pronotum and mesoscutum without or with small punctures which are arranged evenly over the entire surface; mesopleuron projecting ventrally weakly, its lateroventral margin forming an obtuse angle in lateral view (Fig. 19); genal carina bisecting malar space. The larvae develop as parasitoids of crabronid wasps of the subfamily Pemphredoninae. Currently, 26 species are recognised worldwide, most of which are found in the Holarctic Region (Kimsey and Bohart 1991, Wei et al. 2014). A total of eight species are found in Europe (Rosa and Soon 2012), and three are known from the Nordic and Baltic countries (Paukkunen et al. 2014). The status of several European taxa is uncertain, and the genus is in need of taxonomic revision.

Figure 18.

Omalus aeneus ♀. Scale 1 mm.

Key to Omalus species of the Nordic and Baltic countries

1	Mesoscutum wrinkled and dull, without punctures (Fig. 20). Apical notch of T3 deep and triangular (Fig. 21). Body bicoloured with blue, violet or black head and mesosoma, and contrastingly greenish, golden or reddish metasoma	O. biaccinctus (du Buysson)
–	Mesoscutum smooth and shining, without punctures or with evenly distributed punctures (Figs 22, 24). Apical notch smaller (Figs 23, 25). Body concolorous green, blue, violet or mostly black	2
2	Mesoscutum mostly with tiny punctures or impunctate (Fig. 22) and only laterally with short pubescence. If punctures coarser, then flagellomeres slightly longer than broad. Setae laterally on pronotum less than twice as long as diameter of mid-ocellus. Apical notch of T3 shallow (Fig. 23). Flagellomeres as long as or slightly longer than broad	O. aeneus (Fabricius)
–	Mesoscutum with relatively coarse punctures (Fig. 24) and long pubescence. Setae laterally on pronotum at least twice as long as diameter of mid-ocellus. Apical notch of T3 relatively deep (Fig. 25). Flagellomeres short, not longer than broad	O. puncticollis (Mocsáry)

Omalus biaccinctus (du Buysson, 1892)

Figs 20, 21

Ellampus biaccinctus du Buysson (in André), 1892: 152.

Omalus biaccinctus: Trautmann 1927: 41.

Diagnosis

Length 3–5 mm. Both sexes differ from O. aeneus and O. puncticollis by having a bicoloured body (head and mesosoma violet or black, mesosoma reddish or greenish) and a dull and wrinkled mesoscutum without punctures (Fig. 20). The apical notch of T3 is also more deeply triangular (Fig. 21) than in the other two species. The colour of the mesosoma is violet in the female, but dorsally black or dark violet in the male. The metasoma is greenish, golden or reddish in the female, whereas it is greenish, rarely golden or reddish, and dorsally usually black in the male.

Distribution

Denmark, Estonia, Finland, Latvia, Norway, Sweden. Relatively rare. – West Palearctic: from western Europe to western Asia (Linsenmaier 1959).

Biology

Habitat: pine forest margins and clearings, semi-open sandy areas. Occasionally found on flowers of Apiaceae and Asteraceae (Kofler 1975, Rosa 2004). Flight period: June to August. Host: Passaloecus turionum Dahlbom, P. gracilis (Curtis) and P. eremita Kohl (Crabronidae) (Lomholdt 1975, Tormos et al. 1996, Wickl 2001, our own obs.). Adults have been reared from old resin-galls of Retinea resinella (Linnaeus) (Tortricidae) (V. Vikberg, pers. obs.) and pieces of pine wood (Kofler 1975) with host nests inside. The females oviposit in living aphids at the hunting site of their host, and the egg is brought into the host’s nest concealed in the aphid prey (Winterhagen 2015). Thus, the females do not enter the nest of their host for oviposition.

Figures 19–25.

Mesosoma, lateral view (arrow indicating mesopleuron): 19 O. aeneus ♀. Pronotum and mesoscutum, dorsal view: 20 Omalus biaccinctus ♀ 22 O. aeneus ♀ 24 O. puncticollis ♀. T3, postero-dorsal view: 21 O. biaccinctus ♀ 23 O. aeneus ♀ 25 O. puncticollis ♀. Scale 1 mm.

Omalus aeneus (Fabricius, 1787)

Figs 18, 19, 22, 23

Chrysis aenea Fabricius, 1787: 284.

Omalus aeneus: Panzer 1801: 13.

Diagnosis

Length 3–6 mm. The species resembles closely O. puncticollis, but usually has only very small punctures and short pubescence on the mesoscutum (Figs 18, 22), sparser and finer punctation on the pronotum (Figs 18, 22), and a shallower apical notch on T3 (Fig. 23). Some specimens have relatively coarse punctation medially on the pronotum and mesoscutum, but compared to O. puncticollis their pubescence is shorter, flagellomeres are longer, and the apical notch of T3 is shallower. In the female, the body is completely deep blue, violet or green (Fig. 18), whereas in the male it is dorsally black and laterally with green or blue reflections.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Common. – Trans-Palearctic/Holarctic: from western Europe and northern Africa to Japan, China and Taiwan. Possibly accidentally introduced to North America (Kimsey and Bohart 1991) and Australia (Krombein 1979).

Biology

Habitat: forest margins and clearings, semi-open sandy areas. Adults are often observed on sun-exposed leaves of trees and bushes, and they are attracted to honeydew of aphids. Occasionally they are also found on flowers of Apiaceae and Euphorbiaceae (Trautmann 1927, Hoop 1971, Linsenmaier 1997). Flight period: June to August. Host: Passaloecus corniger Shuckard, P. eremita Kohl, P. gracilis (Curtis), P. singularis Dahlbom, P. turionum Dahlbom, Pemphredon lethifer (Shuckard), P. lugubris (Fabricius) and Psenulus pallipes (Panzer) (Crabronidae) (Barbey and Ferriere 1923, Strumia 1997, Gathmann and Tscharntke 1999, our own obs.). The species has been reared from old resin-galls of Retinea resinella (Tortricidae) containing host nests. Females oviposit in live aphids and do not enter the host nest (our own obs.). A similar behaviour has been observed in O. biaccinctus (Winterhagen 2015).

Remarks

Mitochondrial DNA studies indicate that the Nordic and Baltic specimens of O. aeneus belong to at least five genetically distinct lineages (excl. O. puncticollis), and several other lineages have been found in other countries (Paukkunen et al. 2014). It is very likely that more than one species is involved.

Omalus puncticollis (Mocsáry, 1887)

Figs 24, 25

Ellampus puncticollis Mocsáry, 1887: 291.

Omalus puncticollis: Morgan 1984: 16.

Diagnosis

Length 3–6 mm. The species is easily confused with O. aeneus, but the mesoscutum always has relatively large scattered punctures and long setae (Fig. 24). The pronotum has also larger punctures medially (Fig. 24) and the apical notch of T3 is deeper (Fig. 25). The body colouration is similar to O. aeneus. Habitus of large specimens can sometimes resemble small specimens of Pseudomalus violaceus, but the ventral margin of the mesopleuron is not as strongly projecting in O. puncticollis (as in Fig. 19) and the large punctures of the mesoscutum are not clumped postero-medially.

Distribution

Norway, Sweden. Rare. – West Palearctic (?): Europe, Turkey and northern Africa (Linsenmaier 1959, 1968, 1999). The general distribution is poorly known, because many authors have considered O. puncticollis to be conspecific with O. aeneus.

Biology

Habitat: forest margins and clearings, semi-open sandy areas. Adults are usually found sitting on or flying near leaves of trees and bushes, occasionally also on flowers of Apiaceae (Kunz 1994). Flight period: June to August. Host: Passaloecus gracilis (Curtis), P. eremita Kohl, P. corniger Shuckard and P. turionum Dahlbom (Crabronidae) (Spooner 1954, Gauss 1967, Mocsáry 1912).

Remarks

Mitochondrial DNA studies support the recognition of O. puncticollis as a distinct species in relation to O. aeneus (Paukkunen et al. 2014).

Genus Pseudomalus Ashmead, 1902

Figs 2, 26, 27–34

Pseudomalus Ashmead, 1902: 229.

Note

This taxon was raised to generic rank by Kimsey and Bohart (1991). It is characterised by the structure and punctation of the mesosoma: the large punctures are clumped posteriorly between the notauli on the mesoscutum (Fig. 27), and the lateroventral margin of the mesopleuron is strongly projecting ventrally, forming a sharp angle in lateral view (Fig. 28). The posterior margin of T3 is usually deeply notched medially (Figs 31, 32). Pseudomalus is a Holarctic genus with approximately 40 recognised species (Kimsey and Bohart 1991). The larvae are parasitoids of crabronid wasps of the subfamily Pemphredoninae. The European fauna consists of ten species (Rosa and Soon 2012), of which four have been found in the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 26.

Pseudomalus auratus ♀. Scale 1 mm.

Key to Pseudomalus species of the Nordic and Baltic countries

1	Body entirely green to greenish-golden, violet-blue or blackish-green	2
–	Body bicoloured with greenish to bluish head and mesosoma and at least laterally red metasoma	3
2	Body entirely green or green-golden, usually with golden reflections on mesoscutum and mesoscutellum. Metascutellum sharply convex (Fig. 29). Head and mesosoma with short and sparse pubescence, setae not more than twice as long as diameter of mid-ocellus. Smaller species, body length 3–5 mm	P. pusillus (Fabricius)
–	Body uniformly violet-blue or black-green. Metascutellum rounded (Fig. 30). Setae more than twice as long as diameter of mid-ocellus. Larger species, body length 5–7 mm	P. violaceus (Scopoli)
3	Antenna long and slender, medial flagellomeres longer than broad (Fig. 34). Larger species, body length normally not less than 5 mm. Apical notch of T3 triangular, not deeper than broad (Fig. 31)	P. triangulifer (Abeille de Perrin)
–	Antenna shorter, medial flagellomeres approximately as long as broad (Fig. 33). Smaller species, body length usually 3–5 mm, exceptionally up to 6 mm. Apical notch of T3 dorsally arched, as deep as or deeper than broad (Fig. 32)	P. auratus (Linnaeus)

Pseudomalus pusillus (Fabricius, 1804)

Fig. 29

Chrysis pusilla Fabricius, 1804: 176.

Pseudomalus pusillus: Kimsey and Bohart 1991: 268.

Diagnosis

Length 3–5 mm. The species differs from other Pseudomalus species by having an entirely green, green-golden or green-blue body with usually golden reflections on the mesoscutum, mesoscutellum and metanotum. Dark specimens can be confused with unusually dark specimens of P. auratus, but the apex of the metasoma protrudes more narrowly, the metascutellum is more elevated medially (Fig. 29) and the pubescence is shorter.

Distribution

Denmark, Latvia and Lithuania. Very rare. – Trans-Palearctic: from western Europe and northern Africa to Russian Far East (Kurzenko and Lelej 2007).

Biology

Habitat: sparsely vegetated sandy areas, such as river banks and dunes. Adults are attracted to honeydew of aphids (Trautmann 1927) and they are occasionally found on flowers of Apiaceae, Asteraceae, Euphorbiaceae and Resedaceae (Kusdas 1956, Rosa 2004). Flight period: June to August. Host: Passaloecus eremita Kohl, P. insignis (Vander Linden) and Pemphredon lethifer (Shuckard) (Crabronidae) (Benno 1950, Wickl 2001). We consider records mentioning other crabronids (e.g. Rhopalum coarctatum (Scopoli) and species of Trypoxylon Latreille) as hosts to be uncertain, because their biology is quite different from other hosts.

Figures 27–34.

Pronotum and mesoscutum, dorsal view: 27 Pseudomalus auratus ♀. Mesosoma, lateral view (arrow indicating mesopleuron): 28 P. auratus ♀. Mesoscutellum, metanotum and propodeum, lateral view: 29 P. pusillus ♀ 30 P. violaceus ♀. T3, postero-dorsal view: 31 P. triangulifer ♀ 32 P. auratus ♀. Antenna: 33 P. auratus ♀ 34 P. triangulifer ♀. Scale 1 mm.

Pseudomalus auratus (Linnaeus, 1758)

Figs 26, 27, 28, 32, 33

Sphex aurata Linnaeus, 1758: 572.

Pseudomalus auratus: Kimsey and Bohart 1991: 265.

Diagnosis

Length 3–6 mm. Both sexes have a bicoloured body with a blue-green or violet head and mesosoma, and a red (or rarely entirely greenish) metasoma with green reflections (Fig. 26). The species is very similar to P. triangulifer, but the antennal segments are shorter (Fig. 33) and the body is usually smaller. The apical notch of T3 is also deeper and more rounded dorsally (Fig. 32).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Common. – Trans-Palearctic/Holarctic: from western Europe and northern Africa to China, Korea and Japan. Introduced accidentally to North America (Kimsey and Bohart 1991).

Biology

Habitat: forest margins and clearings, gardens and parks. Often found on sun-exposed leaves of deciduous trees and bushes. Adults are attracted to honeydew of aphids and occasionally visit flowers of Apiaceae and Euphorbiaceae (Rosa 2004, our own obs.). Flight period: May to August. Host: cavity-nesting crabronid wasps that prey on aphids, e.g. Passaloecus corniger Shuckard, P. eremita Kohl, P. insignis (Vander Linden), P. gracilis (Curtis), P. monilicornis Dahlbom, P. singularis Dahlbom, P. turionum Dahlbom, Pemphredon inornata Say, P. lethifer (Shuckard), P. lugens Dahlbom, P. lugubris (Fabricius) and P. rugifer (Dahlbom) (Schenck 1856, Benno 1957, van Lith 1958, Brechtel 1986, Blösch 2002, our own obs.), but also Diodontus tristis (Vander Linden), which is a soil-nesting species (Blösch 2002). Host records mentioning other crabronids, such as species of Rhopalum Stephens, Trypoxylon Latreille and Crabro Fabricius, are questionable, because the prey does not consist of aphids in these taxa. Females oviposit in aphids before they have been captured and brought to the nest by the host (our own obs.). A similar behaviour has been observed also in Omalus biaccinctus (Winterhagen 2015) and postulated for Pseudomalus triangulifer (Veenendaal 2011).

Pseudomalus triangulifer (Abeille de Perrin, 1877)

Figs 2, 31, 34

Omalus triangulifer Abeille de Perrin, 1877: 65.

Pseudomalus triangulifer: Kimsey and Bohart 1991: 269.

Diagnosis

Length 6–7 mm. The species resembles closely P. auratus, but the antennal segments are longer (Fig. 34), the body is usually larger and the shape of the apical notch of T3 is shallower and more triangular (Fig. 31). The colour of the metasoma varies from mostly red to almost green. The darkest specimens can be somewhat similar to P. violaceus, but the apical notch is always deeper in P. triangulifer.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – Trans-Palearctic: from Europe and Turkey to China (Linsenmaier 1959, 1968, Rosa et al. 2014).

Biology

Habitat: forest margins and clearings, gardens and parks. Often collected from sun-exposed leaves of trees and bushes. Adults are attracted to honeydew of aphids and occasionally also to flowers of Apiaceae and Euphorbiaceae (Linsenmaier 1997). Flight period: late April to August. Host: Passaloecus insignis (Vander Linden), Pemphredon lugubris (Fabricius), P. lethifer (Shuckard), P. lugens Dahlbom, P. montana Dahlbom and P. rugifer (Dahlbom) (Crabronidae) (Alfken 1915, Strumia 1996, Wickl 2001, Veenendaal 2011, our own obs.). Females probably oviposit in aphids before they have been captured and brought to the nest by the host (Veenendaal 2011).

Pseudomalus violaceus (Scopoli, 1763)

Fig. 30

Sphex violacea Scopoli, 1763: 298.

Chrysis micans Olivier, 1791: 677.

Chrysis fuscipennis Dahlbom, 1829: 15.

Chrysis coerulea Dahlbom, 1831: 33.

Pseudomalus violaceus: Kimsey and Bohart 1991: 270.

Diagnosis

Length 5–8 mm. The species differs from other species of the genus by its completely violet-blue (female) or black-green to black-blue (male) body, and a wide and shallow apical notch on T3. The scapal basin is also higher and dorsally deeply angled. Exceptionally small and worn specimens can be confused with Omalus puncticollis (or O. aeneus), but the mesopleuron of P. violaceus always strongly projects ventrally (as in Fig. 28) and the mesoscutum has large punctures, which are clumped postero-medially (as in Fig. 27).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – Trans-Palearctic (?). Europe, Middle East, Siberia, Manchuria (Linsenmaier 1997). Eastern records could be related to P. bergi Semenov, 1932 (Rosa et al. 2014) or other similar central Asiatic species, e.g. P. bogojavlenskii Semenov, 1932 or P. saturatus Semenov, 1932.

Biology

Habitat: forest margins and clearings. Often found on leaves of sun-exposed deciduous trees and bushes. Flight period: June to August. Adults are attracted to honeydew of aphids (Gauss 1987). Host: Pemphredon lugubris (Fabricius), more rarely also Passaloecus corniger Shuckard and P. eremita Kohl (Crabronidae) (Nielsen 1900, Morgan 1984, Gathmann and Tscharntke 1999, our own obs.). Host records of other species (e.g. Trypoxylon Latreille) are doubtful, because of their deviant biology compared to other hosts. In Finland, the species has been reared from an old gall of Saperda populnea (Linnaeus) (Cerambycidae) on a Populus branch (M. Pentinsaari pers. obs.) and a rotten Alnus stump containing host nests.

Genus Philoctetes Abeille de Perrin, 1879

Figs 35, 36

Philoctetes Abeille de Perrin, 1879: 27.

Note

The taxonomic rank and delineation of Philoctetes has differed among several authors. We follow the definition given by Kimsey and Bohart (1991) and consider it as a distinct genus. Philoctetes is characterised by the following diagnostic features: mesoscutum with large punctures concentrated along notauli; central malar space without carina; mesopleuron rounded and weakly projecting ventrally; metascutellum usually conical; posterior margin of T3 usually deeply notched medially. The hosts consist of crabronid wasps of the subfamily Pemphredoninae. Approximately 40 species are recognised worldwide, and about 30 of these are Palearctic (Kimsey and Bohart 1991). A total of 22 species are known from Europe (Rosa and Soon 2012), but only P. truncatus is found in the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 35.

Philoctetes truncatus ♀. Scale 1 mm.

Philoctetes truncatus (Dahlbom, 1831)

Figs 35, 36

Chrysis truncata Dahlbom, 1831: 35.

Elampus coeruleus of authors, not Dahlbom, 1854.

Philoctetes truncatus: Kimsey and Bohart 1991: 258.

Diagnosis

Length 3–5 mm. The species resembles Omalus aeneus and O. puncticollis by its habitus and colouration. In the female, the body is completely shiny deep blue, violet or green (Fig. 35), whereas it is mainly black with green or blue reflections in the male. Compared to Omalus species, the metascutellum is more sharply elevated and the punctures on the mesoscutum are larger. The apical notch of T3 is shallowly triangular and bordered by a thickened margin (Fig. 36). The lateral margins of T3 are semitransparent and strongly convex adjacent to the apical notch (Fig. 36).

Distribution

Denmark, Estonia, Latvia, Sweden. Rare. New to Latvia (1 ♀, Jekabpils, Avotu iela, 7.–30.VI.2006, leg. P.N. Buhl). – Trans-Palearctic: from western Europe and northern Africa to Russian Far East (Kurzenko and Lelej 2012).

Biology

Habitat: sparsely vegetated sandy areas, sandstone and loess banks (Heinrich 1964, our own obs.). Adults occasionally visit flowers of Apiaceae (Trautmann 1927, Linsenmaier 1997). Flight period: June to July. Host: Diodontus tristis (Vander Linden) (Crabronidae) (Hoop 1961, Linsenmaier 1997, Saure 1998, Jacobs and Kornmilch 2007, our own obs.).

Figure 36.

Philoctetes truncatus ♀ T3, postero-dorsal view. Scale 1 mm.

Genus Elampus Spinola, 1806

Figs 37, 38–46

Elampus Spinola, 1806: 10.

Ellampus Agassiz, 1846: 136.

Notozus Förster, 1853: 351.

Note

This genus has been treated as a subgenus of Omalus by some authors (e.g. Linsenmaier 1959, 1997). It is well characterised by the shape of the metascutellum, which has a large tongue-like projection dorsally (Fig. 38). The posterior margin of T3 is usually extended into a horseshoe-shaped or falcate rim forming truncation (Figs 39–40, 44–46). The female has a row of dense and erect setae along the genal margin (Fig. 41). These setae are replaced by long irregularly placed bristles in the male. The hosts are ground-nesting crabronid wasps, such as Mimesa Shuckard and Mimumesa Malloch (Kimsey and Bohart 1991). The genus is distributed in the Palearctic Region (more than 40 species), North America (8 species), Africa (7 species) and South America (3 species) (Kimsey and Bohart 1991, Linsenmaier 1999, Madl and Rosa 2012). A total of 12 species have been found in Europe (Rosa and Soon 2012), and three of these occur in the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 37.

Elampus panzeri ♀. Scale 1 mm.

Key to Elampus species of the Nordic and Baltic countries

1	Lateral margin of T3 with a narrow notch between apical truncation and semitransparent lateral protrusion (Fig. 39). Apical truncation of T3 horseshoe-shaped with nearly straight ventral margins (Fig. 44)	E. panzeri (Fabricius)
–	Lateral margin of T3 slightly convex or almost straight between apical truncation and semi-transparent lateral protrusion (Fig. 40). Apical truncation with pointed or rounded ventral margins (Figs 45, 46)	2
2	Apical truncation falcate with pointed margins ventrally (Fig. 45). Punctation of T2 dense and regular. Scrobal carina reaches omaulus at its anterior corner (Fig. 42). First flagellomere of female approximately four times as long as broad	E. constrictus (Förster)
–	Apical truncation horseshoe-shaped with rounded margins ventrally (Fig. 46). Punctation of T2 dense, usually with an impunctate central line anteriorly. Scrobal carina reaches omaulus below its anterior protrusion (Fig. 43). First flagellomere of female approximately three times as long as broad	E. foveatus (Mocsáry)

Elampus constrictus (Förster, 1853)

Figs 42, 45

Notozus constrictus Förster, 1853: 336.

Elampus productus of authors, not Dahlbom, 1854.

Ellampus spina of authors, not (Lepeletier, 1806).

Notozus panzeri of authors, not (Fabricius, 1804).

Elampus constrictus: Morgan 1984: 7.

Diagnosis

Length 4–7 mm. The species differs from E. panzeri and E. foveatus by the structure of the apical truncation of T3, which has pointed margins ventrally and resembles a sickle or a boomerang in shape (Fig. 45). Sometimes the apical truncation is very narrow, similar to that of Philoctetes truncatus. The lateral margins of T3 are shallowly concave anterior to the apical truncation. As opposed to E. panzeri and E. foveatus, the scrobal carina reaches the angle of the omaulus (Fig. 42). Both sexes are usually bicoloured with a blue or greenish head and mesosoma, and a reddish metasoma. Entirely greenish or blue specimens are found occasionally.

Distribution

Denmark, Estonia, Finland, Norway, Sweden. Relatively rare. – Trans-Palearctic: widely distributed in the Palearctic Region, from Europe to China (Rosa et al. 2014).

Biology

Habitat: sparsely vegetated sandy areas, heaths. Adults occasionally visit flowers of Apiaceae and Rosaceae (Linsenmaier 1997, Rosa 2004). Flight period: May to July. Host: Mimesa bicolor (Jurine), M. equestris (Fabricius) and M. lutaria (Fabricius) (Crabronidae) (Benno 1950, Lomholdt 1975).

Figures 38–46.

Metascutellum, propodeum and T1, lateral view (arrow indicating metascutellar projection): 38 Elampus foveatus ♀. T3, lateral view: 39 E. panzeri ♀ 40 E. foveatus ♀. Head, lateral view (arrow indicating genal setae): 41 E. panzeri ♀. Mesopleuron, lateral view (arrow indicating junction of omaulus and scrobal carina): 42 E. constrictus ♀ 43 E. foveatus ♀. T3, postero-dorsal view: 44 E. panzeri 45 E. constrictus ♀ 46 E. foveatus ♀. Scale 0.5 mm.

Elampus foveatus (Mocsáry, 1914)

Figs 38, 40, 43, 46

Ellampus foveatus Mocsáry, 1914: 1.

Elampus foveatus: Kimsey and Bohart 1991: 168.

Diagnosis

Length 5–8 mm. The species can be confused with E. constrictus and E. panzeri, but the apical truncation of T3 has rounded margins ventrally and resembles a thick, upside-down U in shape (Fig. 46). The lateral margins of T3 are similar to E. constrictus (Fig. 40). The punctation of T2 is somewhat more irregular than in E. constrictus, and usually an impunctate medial line is formed anteriorly. The scrobal carina ends below the angle of the omaulus (Fig. 43) as in E. panzeri. The head and mesosoma are blue or greenish, and the metasoma is red with green reflections in both sexes.

Distribution

Estonia, Finland, Norway, Sweden. Rare. – Trans-Palearctic: from the Netherlands to Siberia (Usolye-Sibirskoye). The distribution is still poorly known, because many authors have confused E. foveatus with other closely related taxa.

Biology

Habitat: sparsely vegetated sandy areas. In Germany, specimens have been found on Sambucus bushes (Niehuis and Gauss 1996). Flight period: May to July. Host: unknown.

Elampus panzeri (Fabricius, 1804)

Figs 37, 39, 41, 44

Chrysis scutellaris Panzer, 1798: 11, not Fabricius, 1794.

ChrysisPanzeri Fabricius, 1804: 172, replacement name for Chrysis scutellaris Panzer, 1798.

Elampus Panzeri: Latreille 1809: 45.

Notozus constrictus of authors, not Förster, 1853.

Diagnosis

Length 4–8 mm. The species resembles E. constrictus and E. foveatus, but the apical truncation of T3 has angular margins ventrally and resembles a horseshoe in shape (Fig. 44). The lateral margins of T3 also have narrow notches in front of the apical truncation (Fig. 39). The punctation of T2 is somewhat sparser than in E. constrictus and E. foveatus. The scrobal carina is similar to E. foveatus (Fig. 43). Both sexes are bicoloured with a green or blue head and mesosoma, and a red metasoma with green reflections (Fig. 37). Rarely the metasoma can be entirely greenish.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Common. – Trans-Palearctic: Europe, western Asia, Manchuria (Linsenmaier 1959, as E. constrictus).

Biology

Habitat: sparsely vegetated sandy areas, heaths. Adults are occasionally found on flowers of Apiaceae (Heinrich 1964) and stems and inflorescences of grasses (Trautmann 1927, our own obs.). Flight period: late May to August. Adults are also attracted to honeydew of aphids. Host: Mimesa equestris (Fabricius) and M. lutaria (Fabricius) (Crabronidae) (Morice 1903, Benno 1950, our own obs.).

Remarks

The names Elampus constrictus and E. panzeri were erroneously swapped by Trautmann (1927) and later by Linsenmaier (1959, 1997) and other authors. See details from Móczár (1964).

Genus Holopyga Dahlbom, 1845

Figs 3, 7, 47, 48–52

Holopyga Dahlbom, 1845: 4.

Note

This genus consists mainly of broad-bodied wasps, with a body length of 4–9 mm. Morphological characters of the genus include the strongly curved medial vein of the forewing (Fig. 7), the setose medial cell of the forewing, the multidentate tarsal claw (Fig. 3), the carinate and angulate mesopleuron and the evenly rounded posterior margin of T3 (without any distinct notches or prominences). Some species are sexually dimorphic with contrasting colouration in the different sexes (e.g. H. fervida). The biology of most species is poorly known. Apparently, the hosts consist of ground-nesting crabronid and sphecid wasps. Records stating megachilid solitary bees as hosts are questionable due to the lack of supporting data. Holopyga is a large genus with more than 90 recognised species worldwide. The vast majority of these, nearly 70 species, occur in the Palearctic Region (Kimsey and Bohart 1991, Arens 2004). A total of 43 species are known from Europe, and four have been found in the Nordic and Baltic countries (Rosa and Soon 2012, Paukkunen et al. 2014). We have divided the genus into species-groups according to Linsenmaier (1959).

Figure 47.

Holopyga generosa ♂. Scale 1 mm. (Photo: Pekka Malinen)

Key to Holopyga species of the Nordic and Baltic countries

1	Head and mesosoma entirely green-bluish, metasoma dorsally red (Fig. 47)	H. generosa (Förster)
–	Mesosoma dorsally red, or if green-bluish or golden green, then metasoma of the same colour	2
2	Head entirely blue, pronotum, mesoscutum, mesoscutellum and metascutellum dorsally red, rest of mesosoma blue, metasoma dorsally red	H. inflammata (Förster)
–	Head dorsally red, or if green or blue, then whole body green-blue or golden green	3
3	Head and mesosoma partially dorsally red-purple, metasoma red-purple	H. fervida (Fabricius) (female)
–	Body concolorous green-blue or golden greenish	4
4	Punctation of T2 coarse and dense, interstices smaller than puncture diameter (Fig. 49). Medial flagellomeres at most 1.5 times as long as broad (Fig. 51)	H. fervida (Fabricius) (male)
–	Punctation of T2 fine and sparse, interstices larger than puncture diameter at least in the middle (Fig. 50). Medial flagellomeres about 1.5 times as long as broad in the female, and about two times as long as broad in the male (Fig. 52)	H. metallica (Dahlbom)

Holopyga fervida group

Holopyga fervida (Fabricius, 1781)

Figs 49, 51

Chrysis fervida Fabricius, 1781: 456.

Holopyga fervida: Abeille de Perrin 1879: 27.

Diagnosis

Length 4–7 mm. The female and the male are entirely differently coloured. The female is mainly shiny red-purple, but the legs, mesopleuron, metanotum, propodeum, lower part of head and lateral corners of pronotum are blue. The male is entirely green or blue-green, sometimes with golden reflections or a completely golden metasoma. The colouration of the male is similar to H. metallica, but the punctation of the tergites is denser and coarser (Fig. 49) and the antennal segments are shorter (Fig. 51).

Distribution

Denmark. Very rare. Only two records are known from the island of Lolland (1 ♀, Bremersvold, 20.VII.1904, and 1 ♀, Røgebølle, 5.VII.1912, both leg. L. Jørgensen). – West Palearctic: Europe, northern Africa, Turkey, Iran (Kimsey and Bohart 1991, Rosa et al. 2013).

Biology

Habitat: sparsely vegetated sandy areas, loess and clay banks (Kusdas 1956, Hoop 1971, Kunz 1994). Adults visit flowers of Apiaceae, Asteraceae and Euphorbiaceae (Heinrich 1964, Hoop 1971, Linsenmaier 1997, Rosa 2004). Flight period: June to August. Host: unknown.

Figures 48–52.

Head, dorsal view (arrows indicating temples): 48 Holopyga generosa ♀. Metasoma, dorsal view: 49 H. fervida ♂ 50 H. metallica ♂. Antenna: 51 H. fervida ♂ 52 H. metallica ♂. Scale 1 mm.

Holopyga metallica (Dahlbom, 1854)

Figs 50, 52

Hedychrum metallicum Dahlbom, 1854: 68.

Holopyga curvata of authors, not (Förster, 1853).

Holopyga fervida of authors, not (Fabricius, 1781).

Holopyga metallica: Erlandsson 1971: 88.

Diagnosis

Length 5–6 mm. The female is entirely blue or blue-green with golden green reflections on the pronotum, mesoscutum and metasoma. The male is golden green with blue on the metanotum and propodeum. Both sexes resemble the male of H. fervida in colouration, but the punctation of the tergites is finer and sparser (Fig. 50), and the antennal segments are longer in the male (Fig. 52).

Distribution

Finland. Rare. – West Palearctic: only known from Finland and Russian Fennoscandia (Paukkunen et al. 2014).

Biology

Habitat: sparsely vegetated coastal dune areas. Most of the specimens have been collected by sweep nets from grasses and by yellow pan traps. Flight period: June to July. Host: Dryudella stigma (Panzer) (Crabronidae) (according to E. Valkeila’s notes).

Holopyga lucida group (former Holopyga gloriosa group)

Holopyga generosa (Förster, 1853)

Figs 3, 7, 47, 48

Ellampus generosus Förster, 1853: 349.

Holopyga ovata Dahlbom, 1854: 51.

Holopyga amoenula of authors, not Dahlbom, 1845.

Holopyga gloriosa of authors, not (Fabricius, 1793), suppressed name (ICZN 1998).

Holopyga generosa: Linsenmaier 1987: 135.

Holopyga fastuosa ssp. generosa: Linsenmaier 1997: 57.

Diagnosis

Length 7–9 mm. Both sexes are similarly bicoloured with a green or blue head and mesosoma, and a dorsally red metasoma (Fig. 47). The colouration resembles that of Hedychrum gerstaeckeri and the male of H. nobile and H. niemelai, but H. generosa always has multidentate tarsal claws (Fig. 3), angular margins on the head (as in Fig. 48) and an evenly rounded margin of T3 (as in Figs 49, 50). In Hedychrum, the claws are bifid, the head margins are rounded and T3 usually has angular prominences laterally.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Sweden. Common. – Trans-Palearctic: Europe, Asia Minor, northern Africa, China (Linsenmaier 1959).

Biology

Habitat: sparsely vegetated sandy areas, dry meadows. Adults visit flowers of Apiaceae, Asteraceae, Euphorbiaceae, Onagraceae and Rosaceae (Molitor 1935, Linsenmaier 1997, Rosa 2004, our own obs.). Flight period: late May to late August. Host: Astata boops (Schranck) (Crabronidae) (Veenendaal 2012, our own obs.). Females lay their eggs in nymphs of Heteroptera before they have been captured and brought to the nest by the host (Veenendaal 2012).

Holopyga inflammata (Förster, 1853)

Ellampus inflammatus Förster, 1853: 348.

Holopyga gloriosa of authors, not (Fabricius, 1793), suppressed name (ICZN 1998).

Holopyga inflammata: Linsenmaier 1959: 34.

Diagnosis

Length 5–7 mm. Both sexes have similar colouration: the head, propleuron, mesopleuron, propodeum and legs are blue or blue-violet, whereas the pronotum, mesoscutum, mesoscutellum and metascutellum are red. The colouration is relatively similar to the female of H. fervida, but the head is completely blue (without red vertex), the metascutellum is red (not blue) and the mesoscutellum is uniformly punctured (not sparser anteriorly).

Distribution

Finland, Lithuania. Very rare. In Finland, more than 30 specimens were collected in the south-eastern part of the country (Joutseno) in 1957–1960, but currently the species is classified as regionally extinct (Paukkunen 2010). In Lithuania, no records are known since 1970 (Orlovskytė et al. 2010). – West Palearctic: Europe, northern Africa, western Asia (Linsenmaier 1997, 1999).

Biology

Habitat: sparsely vegetated sandy areas. Adults visit flowers of Apiaceae (Brechtel 1985, Rosa 2004). Flight period: early June to early August. Host: unknown.

Genus Hedychrum Latreille, 1802

Figs 4, 53, 54–65

Hedychrum Latreille, 1802: 317.

Note

The genus consists of robust species with a body length ranging from 4 to 10 mm. Characteristic morphological features include the meso- and metatibial pits (Figs 56–58), the enlarged metafemur (Figs 54–55), the apically bifid tarsal claw (Fig. 4) and, in some species the apicomedial tubercle on S4 of the female (Figs 59–60). Many species show sexual dimorphism, in which the mesosoma is strikingly bicoloured in the female (Fig. 47), but uniformly green, blue or violet in the male. The hosts are crabronid wasps of the subfamily Philanthinae. About 150 species are known worldwide, the majority of which occur in the Palearctic Region and Africa (Kimsey and Bohart 1991). The European fauna consists of 17 species and several subspecies, some of which probably would deserve species rank (Rosa and Soon 2012). Five species have been found in the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 53.

Hedychrum niemelai ♀. Scale 1 mm.

Figures 54–65.

Left hindleg, ventral view: 54 Hedychrum nobile ♂ 55 H. nobile ♀. Left mesotibia: 56 H. gerstaeckeri ♂ 57 H. nobile ♂ 58 H. niemelai ♂. S2, ventral view: 59 H. nobile ♀ 60 H. niemelai ♀. T3, dorsal view: 61 H. gerstaeckeri ♂ 62 H. nobile ♂ 63 H. niemelai ♂. Genital capsule, ventral view: 64 H. nobile ♂ 65 H. niemelai ♂, d digitus, c cuspis. Scale 0.5 mm.

Key to Hedychrum species of the Nordic and Baltic countries

1	Male. Anterior surface of metafemur non-metallic black or brown, covered entirely by short adpressed pubescence (Fig. 54). Head and mesosoma dorsally with long and erect pubescence, setae longer than diameter of mid-ocellus	2
–	Female. Anterior surface of metafemur metallic shiny, not entirely covered by short adpressed pubescence (Fig. 55). Head and mesosoma dorsally with short inclined pubescence, setae shorter than diameter of mid-ocellus	6
2	Head and mesosoma dorsally on pronotum, mesoscutum and mesoscutellum with coppery to greenish colour, contrasting with remaining part of the mesosoma, and light brown setae	H. rutilans Dahlbom
–	Head and mesosoma completely green-bluish, with dark setae	3
3	Entire body green-blue	H. chalybaeum Dahlbom
–	Head and mesosoma green-blue, metasoma dorsally metallic red or golden	4
4	Groove on inner surface of mesotibia at least half of tibial length (Fig. 56). Mandible pale brown medially. Punctation of T3 coarse (Fig. 61)	H. gerstaeckeri Chevrier
–	Groove on inner surface of mesotibia shorter or indistinct (Figs 57, 58). Mandible usually not pale brown medially. Punctation of T3 finer (Figs 62, 63)	5
5	Groove on inner surface of mesotibia shallow and narrow, indistinctly delimited (Fig. 57). Punctation of T3 relatively sparse (Fig. 62). Digitus longer than cuspis (Fig. 64)	H. nobile (Scopoli)
–	Groove on inner surface of mesotibia deep and oval, distinctly delimited (Fig. 58). (May be indistinct in small specimens.) Punctation of T3 dense (Fig. 63). Digitus not longer than cuspis (Fig. 65)	H. niemelai Linsenmaier
6	S3 with apicomedial tubercle (Figs 59, 60)	7
–	S3 without apicomedial tubercle	9
7	Head dorsally, pronotum, mesoscutum and mesoscutellum anteriorly bright red	H. chalybaeum Dahlbom
–	Head and mesoscutellum blue-green, pronotum and mesoscutum reddish or golden	8
8	Tubercle of S3 larger, apically slightly rounded, not divided (Fig. 59). Punctation of T3 sparser. Body usually larger, 6–10 mm	H. nobile (Scopoli)
–	Tubercle of S3 smaller, apically divided in the middle (Fig. 60). Punctation of T3 denser. Body usually smaller, 5–8 mm	H. niemelai Linsenmaier
9	Head and mesosoma dorsally with coppery red colour. Head with light brown pubescence	H. rutilans Dahlbom
–	Head and mesosoma entirely blue-green or blue-violet. Head with dark brown pubescence	H. gerstaeckeri Chevrier

Hedychrum gerstaeckeri Chevrier, 1869

Figs 56, 61

Hedychrum Gerstaeckeri Chevrier, 1869: 47.

Diagnosis

Length 4–8 mm. The female differs from the females of other Hedychrum species by having a completely blue, violet-blue or green-blue mesosoma. As in H. rutilans, the female does not have an apicomedial tubercle on S3. Both sexes also have medially pale brown or yellowish mandibles. The male is similar to the female in colouration and can be confused with the males of H. niemelai and H. nobile. However, the punctation of T3 is coarser in H. gerstaeckeri (Fig. 61) and the mesotibial groove is longer and deeper (Fig. 56).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania. Relatively common. – Trans-Palearctic: from western Europe to Japan, China and Taiwan (Rosa et al. 2014).

Biology

Habitat: sparsely vegetated sandy areas, dry meadows. Adults often visit flowers of Apiaceae, Asteraceae and Euphorbiaceae (Trautmann 1927, Kusdas 1956, Kunz 1994, Linsenmaier 1997, Westrich 1979, Rosa 2004, our own obs.). Flight period: mid-June to late August. Host: Cerceris rybyensis (Linnaeus) and C. ruficornis (Fabricius) (Crabronidae) (Berland and Bernard 1938, Grandi 1961, Petit 1975, Westrich 1979, Brechtel 1985, Gayubo et al. 1987, Saure 1998).

Hedychrum rutilans Dahlbom, 1854

Hedychrum rutilans Dahlbom, 1854: 76.

Hedychrum intermedium of authors, not Dahlbom, 1845.

Diagnosis

Length 4–10 mm. The species is usually easy to differentiate from other Hedychrum species by the coppery red colour on the head dorsum, pronotum, mesoscutum and mesoscutellum. Also the pubescence is paler brown than in other species. The ventral part of the head, metanotum, propodeum, mesopleuron and legs are contrastingly blue or blue-green. Sometimes the coppery red colour of the head and/or mesosoma is partially replaced by golden green or blue colour, especially in the male. The mesotibia of the male has a shallow depression on its inner surface, reaching half of the tibial length. The female does not have an apicomedial tubercle on S3.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania. Relatively common. – Trans-Palearctic: Europe, northern Africa, Turkey, southwestern Russia, Siberia (Linsenmaier 1959, 1997, Kimsey and Bohart 1991).

Biology

Habitat: sparsely vegetated sandy areas, dry meadows. Adults often visit flowers of Apiaceae and Asteraceae (Kusdas 1956, Kunz 1994, Rosa 2004, our own obs.). Flight period: early July to late August. Host: Philanthus triangulum (Fabricius) (Crabronidae) (Ferton 1910, Trautmann 1927, Morgan 1984, Veenendaal 1987). The female does not always enter the host nest for ovipositing, but may oviposit on the prey (Apis mellifera Linnaeus) while it is being transported to the nest by the host (Veenendaal 1987, Baumgarten 1995).

Hedychrum nobile (Scopoli, 1763)

Figs 4, 54, 55, 57, 59, 62, 64

Sphex nobilis Scopoli, 1763: 297.

Chrysis lucidula Fabricius, 1775: 358.

Chrysis regia Fabricius, 1793: 243.

Habitat: sparsely vegetated sand and loess areas (Kunz 1994). Adults visit flowers of Apiaceae and Asteraceae (Heinrich 1964). Flight period: July to August. Host: Cerceris interrupta (Panzer) (Crabronidae) (Schmid-Egger 2000). Host records implicating Bembecinus tridens (Fabricius) (Crabronidae: Bembicinae) are probably erroneous, as supporting evidence is lacking.

Genus Hedychridium Abeille de Perrin, 1878

Figs 5, 8, 66, 67–74

Hedychridium Abeille de Perrin, 1878: 3.

Euchrum Semenov, 1954: 103.

Note

This genus comprises a heterogeneous group of small colourful species ranging from 2 to 7 mm in length. Characteristic morphological features include the single perpendicular tooth of the tarsal claw and the transverse pronotal carina (Kimsey and Bohart 1991). The posterior margin of T3 is evenly rounded, without any angular projections. The biology of most species is poorly known, but according to published records, the larvae develop as nest parasites of ground-nesting crabronid wasps and solitary bees. Hedychridium is the second largest genus of Chrysididae and includes more than 300 recognised species worldwide. The highest diversity is found in arid parts of the Holarctic Region and southern Africa. A total of 86 species are known from Europe (Rosa and Soon 2012). Of these, seven occur in the Nordic and Baltic countries (Paukkunen et al. 2014). The genus is here divided into three species-groups according to Linsenmaier (1968, 1997).

Figure 66.

Hedychridium ardens ♀. Scale 1 mm.

Figures 67–74.

Pronotum, mesoscutum and mesoscutellum, dorsal view: 67 Hedychridium ardens ♀, 68 H. coriaceum ♀ 69 H. cupreum ♀ 70 H. purpurascens ♀. Metasoma, dorsal view: 71 H. cupreum ♀ 72 H. purpurascens ♀. Metasoma, ventral view: 73 H. purpurascens ♀. Head, dorsal view (arrows indicating temples): 74 H. roseum ♀. Scale 1 mm.

Key to Hedychridium species of the Nordic and Baltic countries

1	Body entirely green or blue-green. Very small species, body length 2–4 mm	H. zelleri (Dahlbom)
–	Body partially red, golden red or orange. Mostly larger species	2
2	Metasoma non-metallic red or orange, sometimes with weak purple reflections	3
–	Metasoma with strong metallic shine	4
3	Head and pronotum dorsally with coppery red colour. T3 posteriorly with coarse punctures in the male	H. caputaureum Trautmann & Trautmann
–	Head and pronotum entirely blue-green or dark blue. T3 posteriorly with fine punctures in the male	H. roseum (Rossi)
4	Punctation of mesoscutum and mesoscutellum sparse, punctures separated with broad smooth interstices (Figs 69, 70). Metasoma red-purple with blue-green reflections (Figs 71, 72)	5
–	Punctation of mesoscutum and mesoscutellum denser, punctures not separated with broad smooth interstices (Figs 67, 68). Metasoma bright red or golden red with greenish reflections (Fig. 66)	6
5	S2 medially with greenish-golden metallic spot (Fig. 73). Pubescence of body relatively short. Setae on metasoma shorter than the third antennal segment (Fig. 72). Punctation of mesoscutum and mesoscutellum relatively dense (Fig. 70). Scapal basin only medially with fine horizontal ridging	H. purpurascens (Dahlbom)
–	S2 without metallic spot (occasionally with slight metallic sheen). Body with long, erect, whitish pubescence. Setae on metasoma as long as or longer than the third antennal segment (Fig. 71). Punctation of mesoscutum and mesoscutellum sparse (Fig. 69). Scapal basin covered broadly with fine cross-ridging	H. cupreum (Dahlbom)
6	Pronotum and mesoscutum dull with dense coriaceous punctation (Fig. 68). Mesoscutellum with dense punctation and distinct rugae (Fig. 68). Length of malar space less than basal width of mandible	H. coriaceum (Dahlbom)
–	Pronotum and mesoscutum shiny with sparser punctation (Fig. 67). Mesoscutellum with shiny interstices between punctures, without distinct rugae (Fig. 67). Length of malar space equal to basal width of mandible	H. ardens (Coquebert)

Hedychridium monochroum group

Hedychridium zelleri (Dahlbom, 1845)

Hedychrum Zelleri Dahlbom, 1845: 2.

Hedychridium zelleri: du Buysson (in André) 1892: 183.

Diagnosis

Length 2–4 mm. The species differs from other species of Hedychridium by its smaller size and almost completely green or bluish body, which occasionally has weak golden reflections dorsally on the mesosoma. The tarsi are pale brown. Exceptionally small and greenish males of H. ardens can resemble H. zelleri, but they have sparser and coarser punctation dorsally on the mesosoma.

Distribution

Finland. Rare. – West Palearctic: northern and central Europe (Linsenmaier 1959).

Biology

Habitat: sparsely vegetated sandy areas, usually near seashore. Adults are occasionally found on flowers. Flight period: late June to early August. Host: species of Miscophus Jurine (Crabronidae) (Müller 1918, Saure 1998, our own obs.). In central Europe, possibly also Diodontus tristis (Vander Linden) and D. minutus (Fabricius) (Jacobs and Kornmilch 2007).

Hedychridium ardens group

Hedychridium ardens (Coquebert, 1801)

Figs 8, 66, 67

Chrysis ardens Coquebert, 1801: 59.

Hedychrum minutum Lepeletier, 1806: 122.

Chrysis integra Dahlbom, 1829: 17, not Fabricius, 1787.

Hedychridium ardens: Frey-Gessner 1887: 40.

Diagnosis

Length 3–5 mm. Both sexes have coppery red colour dorsally on the head, pronotum, mesoscutum, mesoscutellum and metasoma, whereas the frons, anterior corners of pronotum, metanotum, tibiae and apex of the metasoma are mainly greenish (Fig. 66). The propodeum and the mesopleuron are usually blue. The punctation of the mesoscutum (Fig. 67) is denser than in H. cupreum and H. purpurascens, but not as dense as in H. coriaceum. Small males are sometimes greenish all over with only weak coppery reflections dorsally, and can be confused with H. zelleri. The punctation of the mesoscutum is however sparser and coarser than in H. zelleri.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Common. – Trans-Palearctic: Europe, Mongolia, Russian Far East (Kimsey and Bohart 1991, Linsenmaier 1997, Kurzenko and Lelej 2007).

Biology

Habitat: sparsely vegetated sandy areas, dunes, dry meadows. Adults visit flowers of Apiaceae, Asteraceae, Crassulaceae, Euphorbiaceae and Rosaceae (Molitor 1935, our own obs.). Flight period: late May to late August. Host: Diodontus tristis (Vander Linden), Oxybelus bipunctatus Olivier, Tachysphex nitidus (Spinola), T. obscuripennis (Schenck) and T. pompiliformis (Panzer) (Crabronidae) (Trautmann 1927, Berland and Bernard 1938, Benno 1950, Else 1973, Kofler 1975, Morgan 1984, van der Smissen 2001).

Hedychridium coriaceum (Dahlbom, 1854)

Fig. 68

Hedychrum coriaceum Dahlbom, 1854: 88.

Hedychridium coriaceum: du Buysson (in André) 1892: 195.

Diagnosis

Length 3–5 mm. The species is characterised by the very dense and fine punctation of the pronotum and mesoscutum, whereby the surface appears completely dull (Fig. 68). The mesoscutellum also has dense punctation and rugae between the punctures (Fig. 68). In H. ardens, H. cupreum and H. purpurascens the punctation is sparser and the surface shinier. The colour of the vertex, pronotum, mesoscutum and mesoscutellum is brownish red (Fig. 68). The frons, anterior corners of pronotum, metanotum, propodeum, mesopleuron and tibiae are mainly green or blue. The metasoma is coppery red or sometimes greenish.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Sweden. Relatively rare. – Trans-Palearctic: from Europe and northern Africa to China (Linsenmaier 1959, Rosa et al. 2014). (In northern Africa represented by ssp. jendoubense Linsenmaier, 1987).

Biology

Habitat: sparsely vegetated sandy areas. Adults visit flowers of Asteraceae, Euphorbiaceae and Rosaceae (Heinrich 1964, Hoop 1971, Linsenmaier 1997, Rosa 2004, our own obs.). Flight period: early June to late August. Host: Lindenius albilabris (Fabricius) (Crabronidae) (Arnold 1908, 1910, Mortimer 1913, Morgan 1984, Tischendorf 1998) and possibly also Oxybelus uniglumis (Linnaeus) (Alfken 1915).

Hedychridium cupreum (Dahlbom, 1845)

Figs 69, 71

Hedychrum cupreum Dahlbom, 1845: 3.

Remarks

H. caputaureum was recently considered to be a central and northern European subspecies of H. chloropygum (Arens 2010). However, as these taxa are both well characterised and have partially overlapping distribution areas, we treat them as separate species following Paukkunen et al. (2014).

Hedychridium roseum (Rossi, 1790)

Figs 5, 74

Chrysis carnea var. rosea Rossi, 1790: 75.

Chrysis rufa Panzer, 1800: 16.

Chrysis Rosae Dahlbom, 1829: 13.

Hedychridium roseum: Abeille de Perrin 1879: 35.

Diagnosis

Length 5–8 mm. The species differs from H. caputaureum by having the vertex (Fig. 74) and mesosoma completely blue or greenish, sometimes nearly black, without any trace of coppery red colour. Some males, however, have golden reflections on the pronotum and mesoscutum. The punctation of the tergites is finer and denser throughout, not becoming coarser posteriorly on T2 and T3 as in most males of H. caputaureum. The metasoma is similarly non-metallic red or orange as in H. caputaureum, and may have weak metallic violet reflections posteriorly.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Common. – Trans-Palearctic: from western Europe to Siberia, China and Russian Far East (Linsenmaier 1959, Kurzenko and Lelej 2007, Rosa et al. 2014).

Biology

Habitat: sparsely vegetated sandy areas, dry meadows. Adults visit flowers of Apiaceae, Asteraceae, Crassulaceae and Euphorbiaceae (Trautmann 1927, Kusdas 1956, Heinrich 1964, Brechtel 1985, Linsenmaier 1997, Rosa 2004). Flight period: late June to late August. Host: primarily Astata boops (Schranck) (Crabronidae) (Reder 2010, our own obs.), but possibly also Dryudella stigma (Panzer), Tachysphex pompiliformis (Panzer) and Harpactus tumidus (Panzer) (Crabronidae) (Shuckard 1837, Müller 1918, Forsius 1925, Doronin 1996, Saure 1998).

Tribe Chrysidini

Members of this tribe are characterised by the simple untoothed tarsal claw, the transverse subapical pit row on T3, and the transverse preoccipital welt or carina (Kimsey and Bohart 1991). With more than 1.500 species and 30 genera, Chrysidini is the largest tribe of Chrysididae. A total of 11 genera are known from Europe (Rosa and Soon 2012), five of which are found in the Nordic and Baltic countries (Paukkunen et al. 2014).

Genus Pseudospinolia Linsenmaier, 1951

Figs 9, 75

Pseudospinolia Linsenmaier, 1951: 31.

Note

This taxon has been treated as a subgenus of Euchroeus Latreille or Spinolia Dahlbom by some authors (Linsenmaier 1959, Morgan 1984, Arens 2014). We follow the classification of Kimsey and Bohart (1991) and regard it as a distinct genus. Characteristic morphological features include the broadly open marginal cell (Fig. 9), the edentate T3, the deep and subdivided depression laterally on the pronotum, and the elongate transverse frontal carina connecting the compound eyes. Members of the genus parasitise solitary vespids of the subfamily Eumeninae (e.g. Odynerus Latreille). Approximately 15 species are recognised worldwide, most of which occur in the Palearctic Region, particularly in the Middle East and the Mediterranean region (Kimsey and Bohart 1991). The European fauna consists of eight species (Rosa and Soon 2012), and one, P. neglecta, is found in the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 75.

Pseudospinolia neglecta ♀. Scale 1 mm.

Pseudospinolia neglecta (Shuckard, 1837)

Figs 9, 75

Chrysis austriaca Dahlbom, 1829: 14, not Fabricius, 1804.

Chrysis neglecta Shuckard, 1837: 169.

Chrysis integrella Dahlbom, 1854: 133.

Pseudospinolia neglecta: Kimsey and Bohart 1991: 548.

Diagnosis

Length 5–9 mm. Both sexes are bicoloured with a green or blue head and mesosoma, and a golden red metasoma (Fig. 75). The vertex and the anterior margin of the pronotum often have golden reflections. The metasoma is very finely and densely punctured on the tergites, causing the surface to appear dull (Fig. 75). The posterior margin of T3 is edentate. The species can be confused with similarly coloured species of Chrysura, but the radial sector vein of the forewing does not reach the wing margin (Fig. 10).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – Trans-Palearctic/Holarctic? Europe, Asia, Russian Far East, China, USA, Canada (Kimsey and Bohart 1991, Kurzenko and Lelej 2007, Rosa et al. 2014). Possibly accidentally introduced to North America (Bohart and Kimsey 1982).

Biology

Habitat: sparsely vegetated areas with clay or sandy soil, gardens with clay structures, such as barn walls. Adults occasionally visit flowers of Asteraceae, Crassulaceae and Rosaceae (Kusdas 1956, Heinrich 1964, Ressl 1966, Linsenmaier 1997, Rosa 2004, 2006). Flight period: late May to mid-August. Host: Odynerus spinipes (Linnaeus) and O. reniformis (Gmelin) (Vespidae) (Smith 1862, Adlerz 1910, Trautmann 1927, Linsenmaier 1959, Banaszak 1980, Morgan 1984), possibly also Ancistrocerus parietum (Linnaeus) and Gymnomerus laevipes (Shuckard) (Dahlbom 1854, Berland and Bernard 1938). Host records mentioning bees of the genera Osmia Panzer and Heriades Spinola (Megachilidae) are doubtful, as noted by Kunz (1994).

Genus Spinolia Dahlbom, 1854

Fig. 76

Spinolia Dahlbom, 1854: 363.

Achrysis Semenov, 1892: 486.

Note

Members of this genus are characterised by the broadly open marginal cell, the U-shaped projection on the lower mesopleuron, the dentate segments of the ovipositor, and the frons with two rounded, flattened and usually striate areas particularly in the male (Kimsey and Bohart 1991). The medial cell of the forewing is asetose. The larvae are nest parasites of solitary vespids of the subfamily Eumeninae (e.g. Pterocheilus Klug and Hemipterochilus Ferton). The genus consists of 15 Palearctic species, most of which are found in northern Africa, Middle East and central Asia. In Europe, seven species are known (Rosa and Soon 2012), of which one, S. unicolor, is recorded from the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 76.

Spinolia unicolor ♀. Scale 1 mm.

Spinolia unicolor (Dahlbom, 1831)

Figs 76

Chrysis unicolor Dahlbom, 1831: 32.

Spinolia unicolor: du Buysson (in André) 1893: 244.

Diagnosis

Length 4–6 mm. The species is characterised by the entirely blue, greenish or violet-blue body, and the forewing radial sector vein, which ends remote from the wing margin (Fig. 76). The T3 is posteriorly edentate and has a small tooth anteriorly.

Distribution

Denmark, Latvia, Sweden. Very rare. – Trans-Palearctic: from western Europe to Mongolia (Linsenmaier 1959).

Biology

Habitat: xerothermic sparsely vegetated sandy areas, often close to the seashore. Adults occasionally visit flowers of Asteraceae, Lamiaceae and Rosaceae (Trautmann 1927, Benno 1950, Linsenmaier 1959). Flight period: late June to mid-August. Host: Pterocheilus phaleratus (Panzer) (Vespidae: Eumeninae) (Erlandsson 1968, Sörensson and Cederberg 2010).

Genus Chrysis Linnaeus, 1761

Figs 6, 10, 77–92, 93–101, 102–109, 110–125, 126–137, 138–144, 145–149, 150–164, 165–178, 179, 180–186, 187–195

Chrysis Linnaeus, 1761: 414.

Chrysogona Förster, 1853: 327.

Tetrachrysis Lichtenstein, 1876: 27.

Hexachrysis Lichtenstein, 1876: 27.

Note

With more than a thousand currently recognised species, Chrysis is the largest and most heterogeneous genus of Chrysididae. It is best defined by a combination of several variable and non-unique characters, such as the closed or nearly closed forewing marginal cell, the usually four- or six-toothed posterior margin of T3, and the usually distinct transverse frontal carina on the frons. Members of the genus parasitise a wide range of solitary wasps and bees in the families Vespidae, Sphecidae, Crabronidae, Megachilidae and Apidae. They are found worldwide, but the vast majority of species is found in the Holarctic and Afrotropical Regions. The European fauna consists of nearly 190 species and numerous subspecies (Rosa and Soon 2012). Up to now, 35 species have been found in the Nordic and Baltic countries (Paukkunen et al. 2014). The genus was first formally divided into species-groups by Linsenmaier (1959). Our classification of the species-groups follows Kimsey and Bohart (1991).

Figures 77–92.

T3, dorsal view: 77 Chrysis succincta ♀ 78 C. illigeri ♀ 79 C. viridula ♀ 80 C. equestris ♀ 81 C. sexdentata ♀ 82 C. indigotea ♂ 83 C. iris ♂ 84 C. impressa ♀ 85 C. impressa ♂ 86 C. angustula ♂ 87 C. subcoriacea ♂ 88 C. longula ♂. Ovipositor, dorsal view: 89 C. equestris ♀ 90 C. zetterstedti ♀ 91 C. solida ♀ 92 C. impressa ♀. Scale 1 mm.

Figures 93–101.

Metasoma, dorsal view: 93 Chrysis fulgida ♂ 94 C. rutilans ♀ 95 C. splendidula ♀ 96 C. ruddii ♀ 97 C. vanlithi ♀ 98 C. clarinicollis ♀ 99 C. angustula ♀ 100 C. mediata ♀ 101 C. solida ♀. Scale 1 mm.

Figures 102–109.

Metasoma, dorsal view: 102 C. longula ♀ 103 C. corusca ♀ 104 C. ignita ♀ 105 C. ignita ♂ 106 C. borealis sp. n. ♀ 107 C. impressa ♂. Metasoma, lateral view: 108 C. impressa ♂ 109 C. subcoriacea ♀. Scale 1 mm.

Figures 110–125.

Metasoma, ventral view: 110 Chrysis bicolor ♀, 111 C. illigeri ♀ 112 C. equestris ♀ 113 C. zetterstedti ♀ 114 C. splendidula ♀ 115 C. vanlithi ♀ 116 C. impressa ♀ 117 C. subcoriacea ♀ 118 C. leptomandibularis ♀ 119 C. angustula ♀ 120 C. solida ♀ 121 C. longula ♀ 122 C. ignita ♀ 123 C. corusca ♀ 124 C. borealis sp. n. ♀ 125 C. solida ♂. Scale 1 mm.

Figures 126–137.

Metasoma, ventral view: 126 Chrysis angustula ♂ 127 C. leptomandibularis ♂ 128 C. corusca ♂ 129 C. subcoriacea ♂ 130 C. ignita ♂ 131 C. schencki ♂ 132 C. impressa ♂ 133 C. borealis sp. n. ♂. Genital capsule, dorsal view: 134 C. equestris ♂ 135 C. zetterstedti ♂ 136 C. borealis sp. n. ♂ 137 C. solida ♂. Scale 1 mm (Figs 126–133) and 0.5 mm (Figs 134–137).

Figures 138–144.

Head, lateral view: 138 Chrysis bicolor ♀ 139 C. illigeri ♀ 140 C. vanlithi ♀ 141 C. impressa ♀ 142 C. leptomandibularis ♀ 143 C. schencki ♀ 144 C. corusca ♀. Scale 1 mm.

Figures 145–149.

Head, lateral view: 145 C. leptomandibularis ♂ 146 C. subcoriacea ♂ 147 C. ignita ♂ 148 C. schencki ♂ 149 C. impressa ♂. Scale 1 mm.

Figures 150–164.

Head, frontal view (arrow indicating frontal carina): 150 Chrysis leachii ♀ 151 C. equestris ♀ 152 C. zetterstedti ♀ 153 C. terminata ♀. Head, dorsal view (arrow indicating frontal carina): 154 C. terminata ♀. Mandible (arrow indicating subapical tooth): 155 C. brevitarsis ♀. Pronotum and mesoscutum, dorsal view: 156 C. indigotea ♀, 157 C. brevitarsis ♀ 158 C. borealis sp. n. ♀. Pronotum, dorsal view: 159 C. vanlithi ♀ 160 C. subcoriacea ♂ 161 C. longula ♂. Pronotum and mesoscutum, dorsal view (arrow indicating lateral field of mesoscutum): 162 C. clarinicollis ♀ 163 C. ignita ♀ 164 C. impressa ♀. Scale 1 mm.

Key to Chrysis species of the Nordic and Baltic countries

1	Posterior margin of T3 medially pointed or rounded without teeth (Fig. 77)	2
–	Posterior margin of T3 with distinct teeth or angular prominences (Figs 78–88)	4
2	Posterior margin of T3 almost evenly rounded, medially not pointed. Mesoscutum medially dark blue or blackish and laterally green or blue with golden reflections	C. gracillima Förster
–	Posterior margin of T3 medially somewhat pointed (Fig. 77). Mesoscutum completely red or golden greenish, medially not darker than laterally	3
3	Scapal basin largely smooth (female) or finely punctured (male) without fine cross-ridging. Mesoscutellum blue. Larger species, 4–8 mm	C. succincta Linnaeus
–	Scapal basin medially with fine cross-ridging in both sexes (Fig. 150). Mesoscutellum golden red or greenish-golden. Smaller species, 3–6 mm	C. leachii Shuckard
4	Medial teeth on the posterior margin of T3 extending distinctly further posteriorly than lateral teeth, and usually located in close proximity (Fig. 78)	5
–	Medial teeth on the posterior margin of T3 not extending distinctly further posteriorly than lateral teeth, and/or not located in close proximity (Fig. 79–88)	7
5	Mesoscutum blue or blackish. Second metatarsomere at least 3.5 times as long as broad (Fig. 171)	C. westerlundi Trautmann
–	Mesoscutum golden red or golden green. Second metatarsomere at most 2.5 times as long as broad	6
6	Malar space shorter, at most 0.75 times basal width of mandible (Fig. 138). Punctation of T2 dense. Metascutellum raised medially. Black spots of S2 strongly oblique posteriorly in female (Fig. 110)	C. bicolor Lepeletier
–	Malar space longer, in profile equal to basal width of mandible (Fig. 139). Punctation of T2 sparser. Metascutellum flat medially. Black spots of S2 not strongly oblique posteriorly in female (Fig. 111)	C. illigeri Wesmael
7	Posterior margin of T3 with six teeth (Figs 80–81)	8
–	Posterior margin of T3 with four teeth, these sometimes only shallow projections (Figs 79, 82–88)	10
8	Metasoma dorsally purple red. Lateral teeth of T3 sharp (Fig. 81)	C. sexdentata Christ
–	Metasoma multicoloured, T1 and T2 dark blue or black with golden green or golden red bands posteriorly. Lateral teeth of T3 rounded (Fig. 80)	9
9	Black spots of S2 fused and short, not or only slightly extending to lateral margins of sternite (Fig. 112). Ovipositor relatively broad and strongly chitinised (Fig. 89). T5 of female with transverse striae and longitudinal medial groove (Fig. 89). Head distinctly wider than high (Fig. 151). Male genitalia with broad notch between parameres, gonostyle very short and cuspis apically curved (Fig. 134). Propodeal tooth slightly convex or straight ventrally	C. equestris Dahlbom
–	Black spots of S2 fused and broad, widely extending to the lateral margins of the sternite (Fig. 113). Ovipositor narrow and weakly chitinised (Fig. 90). T5 of female without transverse striae and medial groove (Fig. 90). Head only slightly wider than high (Fig. 152). Male genitalia with narrow notch between parameres, gonostyle apically elongated (as long as cuspis) and cuspis apically straight (Fig. 135). Propodeal tooth weakly lobate ventrally	C. zetterstedti Dahlbom
10	Metasoma completely blue or blue-green, without red colour	11
–	Metasoma not completely blue or blue-green, always with red or golden colour	12
11	Mesoscutum medially not darker than laterally. Black spots of S2 large and long, almost extending to middle of sternite. T3 of female with distinct transverse bulge anterior to pit row. Male with short triangular apical teeth separated by wide intervals (Fig. 83). Ovipositor narrow (as in Fig. 92)	C. iris Christ
–	Mesoscutum distinctly darker medially than laterally (Fig. 156). Black spots of S2 smaller, distinctly separated in the middle. T3 of female without transverse bulge anterior to pit row. Male with spiniform apical teeth separated by narrow intervals (Fig. 82). Ovipositor broad (as in Fig. 91)	C. indigotea Dufour & Perris
12	T1 blue, T2 entirely golden red (female) or with large dark blue dorsal spot (male) (Fig. 93)	C. fulgida Linnaeus
–	Both T1 and T2 golden red or reddish	13
13	T3 completely green, blue or violet, in contrast with colour of T1 and T2	14
–	T3 golden red or reddish, at most with blue or violet apical rim	17
14	Mesosoma dorsally bright red	15
–	Mesosoma dorsally blue-green or blue-violet	16
15	Setae medially on metatibia longer than width of tibia (Fig. 166). Head completely blue-green or blue-violet. Mesoscutum completely red in male	C. viridula Linnaeus
–	Setae medially on metatibia shorter than width of tibia (Fig. 165). Head dorsally red in female. Mesoscutum medially green or blue in male	C. pulcherrima Lepeletier
16	Body slender, metasoma narrower (Fig. 94). T2 without or with very weak longitudinal keel medially (Fig. 94), posteriorly without lifted margin. Punctation of T2 relatively fine and sparse, interstices as large as or larger than puncture diameter (Fig. 94). Head in frontal view as wide as high. Black spots of S2 three quarters length of sternite	C. rutilans Olivier
–	Body robust, metasoma broader (Fig. 95). T2 with a distinct smooth longitudinal keel medially (Fig. 95), posteriorly with slightly raised margin. Punctation of T2 relatively coarse and dense, interstices smaller than puncture diameter (Fig. 95). Head in frontal view slightly wider than high. Black spots of S2 half of length of sternite (Fig. 114)	C. splendidula Rossi
17	Apical rim of T3 blue or violet, remaining tergite red	18
–	Apical rim of T3 golden red, of same colour as remaining tergite	19
18	Meso- and metascutellum golden red	C. scutellaris Fabricius
–	Mesosoma entirely blue-violet	C. graelsii Guérin-Méneville
19	Spurs of metatibia approximately equal in length (Fig. 167). Female metatarsus shorter than metatibia, second tarsomere twice as long as broad in lateral view (Fig. 169). Mandible thicker (in male, medial width of mandible about two thirds of its basal width; in female, medial width of mandible more than half of its basal width), with or without subapical tooth. Flagellomeres distinctly nodular in female (Fig. 172)	20
–	Spurs of metatibia distinctly unequal in length (Fig. 168). Female metatarsus longer than metatibia, second tarsomere at least three times longer than broad in lateral view (Fig. 170). Mandible thinner (in male, medial width of mandible less than two thirds of its basal width; in female, medial width of mandible not more than half of its basal width), always without subapical tooth. Flagellomeres not distinctly nodular in female	21
20	Mandible with subapical tooth (Fig. 155). Female mesoscutum laterally with scattered punctation (Fig. 157)	C. brevitarsis Thomson
–	Mandible without subapical tooth. Female mesoscutum laterally with dense punctation	C. pseudobrevitarsis Linsenmaier
21	Pronotum short, length less than one fourth of its width (Fig. 159). Malar space long, approximately as long as broad in female (Fig. 140), and somewhat shorter in male. F1 of antenna without metallic sheen. Frons with dense adpressed white pubescence	22
–	Pronotum longer, length at least one fourth of its width (Figs 158, 160–164). Malar space short, shorter than broad in both female (Figs 141–144) and male (Figs 145–149). F1 with metallic sheen (often very weak). Frons usually with sparser and more erect pubescence	23
22	Punctation of tergites very fine and dense throughout, punctures of uniform size, surface dull (Fig. 96). Sternites and legs ventrally coppery red	C. ruddii Shuckard
–	Punctation of tergites coarser and sparser, punctures of variable size, surface shiny (Fig. 97). Sternites and legs ventrally always greenish (Fig. 115)	C. vanlithi Linsenmaier
23	Frontal carina with four tooth-like tubercles medially (Figs 153, 154). Punctation of tergites coarse throughout (as in Figs 104, 105). Head and mesosoma dorsally with white pubescence. Apical teeth of T3 relatively long and sharp (as in Figs 104, 105)	C. terminata Dahlbom
–	Frontal carina without four tooth-like tubercles. Punctation of tergites, colour of pubescence and shape of apical teeth variable	24
24	Pronotum and mesoscutellum uniformly greenish (Fig. 162). T1 anteriorly and laterally blue-green or green, dorsally golden red (Fig. 98). Shape of metasoma broad and compact (Fig. 98). Apical rim short and intervals between apical teeth shallow (Fig. 98)	C. clarinicollis Linsenmaier
–	Pronotum blue or violet framed by lighter colour, mesoscutellum medially darker than laterally (Figs 158, 160, 161, 163, 164). T1 laterally golden red or only slightly greenish, dorsally golden red. Shape of metasoma, apical rim and apical teeth variable (Figs 99–107)	25
25	Female. With ovipositor. Posterior margin of S4 opaque and angled (Figs 116–124). T3 dorsally straight or concave in lateral view (Fig. 109)	26
–	Male. Without ovipositor. Posterior margin of S4 semitransparent and almost straight (Figs 125–133). T3 dorsally convex in lateral view (Fig. 108)	36
26	T2 and T3 laterally completely dull, with dense coriaceous microsculpture between punctures (Fig. 109). Elongate and usually large species, ovipositor thin (as in Fig. 92), S2 red with long black spots (Fig. 117)	C. subcoriacea Linsenmaier
–	T2 and T3 laterally with shiny surface between punctures (as in Fig. 108). Body shape, breadth of ovipositor and colouration of S2 variable	27
27	Mandible extremely thin (in lateral view, medially as broad as apical segment of labial palp) (Fig. 142), dorsally smooth and impunctate. Body slender and elongate, metasoma with almost parallel sides (as in Fig. 99). S2 greenish with short, rounded black spots (Fig. 118). Mesoscutum laterally with wide, strongly shining interstices between punctures	C. leptomandibularis Niehuis
–	Mandible thicker (in lateral view, medially broader than apical segment of labial palp), dorsally always punctate. Shape of body, colouration of S2 and punctation of mesoscutum variable	28
28	Body slender and elongate, metasoma with almost parallel sides (Fig. 99). Punctation of T2 anteriorly finer than on T1, posteriorly very sparse (Fig. 99). T3 long, deeply depressed medially and strongly shining. Apical teeth short and blunt, with a wide and shallow central interval (Fig. 99). S2 with rectangular black spots and golden red colour (Fig. 119). Ovipositor narrow (as in Fig. 92)	C. angustula Schenck
–	Body not as slender and elongate (Figs 100–104, 106). Punctation of T2 variable (Figs 100–104, 106). T3 shorter, more shallowly depressed medially and not as strongly shining. Apical teeth longer and/or sharper, shape of central interval variable (Figs 100–104, 106). Black spots of S2 not as distinctly rectangular, colouration variable (Figs 116, 120–124). Ovipositor narrow (Fig. 92) or broad (Fig. 91)	29
29	Ovipositor broad (T5 broader than long) (Fig. 91), often only its apex exerted (Fig. 100). S2 blue-green with relatively short and rounded black spots (Fig. 120). Mandible thick (in lateral view, medial width of mandible about half of its basal width) (as in Fig. 144). Apical rim of T3 long and central interval between apical teeth often angular	30
–	Ovipositor narrow (T5 longer than broad) (Fig. 92). Colouration of S2 (Figs 116, 121–124) and thickness of mandible variable (Figs 141, 143, 144). Apical rim usually shorter and central interval more widely arcuate	31
30	Metasoma broader, with slightly convex sides (Fig. 100). Punctation of T2 anteriorly sparser and finer, with shining interstices (Fig. 100). T3 with slightly sparser punctation, its surface thereby shinier (Fig. 100). Head narrower, in frontal view only slightly broader than high. Colour of mesosoma lighter and more greenish. Usually larger species. Hosts soil-nesting species of Odynerus	C. mediata Linsenmaier
–	Metasoma more elongate, with more parallel sides (Fig. 101). Punctation of T2 anteriorly denser and coarser, without shining interstices (Fig. 101). T3 with dense punctation, its surface dull (Fig. 101). Head broader, in frontal view distinctly broader than high. Colour of mesosoma darker and more bluish. Usually smaller species. Hosts cavity-nesting species of Ancistrocerus and Euodynerus	C. solida Haupt
31	Metasoma elongate with almost parallel sides (Figs 102, 103). S2 with long and narrow black spots (Figs 121, 123). Mandible thicker (medial width of mandible about half of its basal width) (Fig. 144)	32
–	Metasoma with more convex sides (Figs 104, 106). S2 with shorter black spots (Figs 116, 122, 124). Mandible thinner (medial width of mandible less than half of its basal width) (Figs 141, 143)	33
32	T2 anteriorly with dense, deep and coarse punctation, punctures becoming much sparser and finer posteriorly (Fig. 102). T3 shiny, without microsculpture (Fig. 102). S2 red (Fig. 121).Mandible longer and thinner. Metascutellum medially flat. Usually larger species	C. longula Abeille de Perrin
–	T2 anteriorly with somewhat sparser and finer punctation, punctures becoming slightly sparser and finer posteriorly (Fig. 103). T3 shiny or with weak microsculpture (Fig. 103). S2 greenish (Fig. 123). Mandible shorter and thicker. Metascutellum medially more convex. Usually smaller species	C. corusca Valkeila
33	Apical teeth of T3 sharply produced and apical rim with almost parallel lateral margins (Fig. 104). Punctation of T2 and T3 coarse throughout (Fig. 104). Pubescence of vertex whitish. S2 green or blue with rectangular black spots (Fig. 122). Mesoscutum shiny blue or greenish, punctures of same colour as interstices (Fig. 163). Medial furrow of pronotum narrow (Fig. 163)	C. ignita (Linnaeus)
–	Apical teeth of T3 not as sharply produced and apical rim with more angled lateral margins (Figs 84, 106). Punctation of T2 and T3 finer (Figs 84, 106). Pubescence of vertex whitish or brownish. Colouration of S2 (Figs 116, 124) and mesoscutum (Figs 158, 164) variable. Medial furrow of pronotum broader (Figs 158, 164)	34
34	Mandible very thin (medial width of mandible not more than one third of its basal width), basally strongly narrowing in lateral view (Fig. 143). Scapal basin with sparse and well defined punctation. Vertex with light brown pubescence. T3 relatively dull with distinct microsculpture between punctures. Mesoscutum dark blue to almost black. Metasoma usually more elongate	C. schencki Linsenmaier
–	Mandible thicker (medial width of mandible more than one third of its basal width), gradually narrowing towards the apex (Fig. 141). Scapal basin with denser and more coriaceous punctation. Vertex with brown or whitish pubescence. T3 usually shinier. Colour of mesoscutum variable (Figs 164, 158, 179). Metasoma usually broader and more compact	35
35	Mesoscutum laterally with green or blue punctures and black interstices (Fig. 164). Punctures relatively large (Fig. 164). F1 1.3–1.5 times as long as F2. S2 posteriorly usually red with relatively rounded black spots (Fig. 116). Pubescence of vertex brownish	C. impressa Schenck
–	Mesoscutum entirely blue, violet or black, punctures and interstices generally of same colour (Figs 158, 179). Punctures smaller (Figs 158, 179). F1 1.5–1.7 times as long as F2. S2 dark green or blue with almost rectangular black spots (Fig. 124). Pubescence of vertex whitish or light brown	Chrysis borealis sp. n.
36	(Males.) Mandible thin (medial width of mandible about one third of its basal width) (Fig. 145), dorsally smooth, without or with only barely visible punctures. Body small and slender, metasoma elongate with almost parallel sides (habitus similar to C. angustula). S2 greenish with short rounded black spots (Fig. 127). Posterior margin of propodeal tooth virtually straight and perpendicular to body axis	C. leptomandibularis Niehuis
–	Mandible thicker (medial width of mandible more than one third of its basal width), its dorsal surface always with small punctures. Body size and shape variable. Colouration of S2 variable (Figs 125, 126, 128–133). Posterior margin of propodeal tooth straight, convex or concave	37
37	Colour of S2 golden or reddish, with long rectangular black spots (Fig. 126). Body slender, metasoma with approximately parallel sides (habitus similar to C. leptomandibularis). Punctation of T2 anteriorly usually finer than on T1, becoming sparser posteriorly, surface thereby strongly shining. T3 shiny, transition of lateral margin into lateral apical teeth straight, apical rim and teeth short (Fig. 86)	C. angustula Schenck
–	Colour of S2 variable, black spots usually not rectangular (Figs 125, 128–133). Body usually more robust, metasoma with more convex sides. Punctation of T2 anteriorly slightly finer or as coarse as on T1, surface posteriorly not as shining. T3 shiny or dull, transition of lateral margin into lateral apical teeth straight or concave, apical rim and teeth longer (Figs 85, 87–88)	38
38	Inner margin of paramere angled (Fig. 137). Punctation of T2 anteriorly usually finer than on T1. T3 with very fine and regular punctation. S2 usually greenish with relatively short black spots (Fig. 125). Mandible relatively thick (in lateral view, medial width of mandible more than half of its basal width)	39
–	Inner margin of paramere rounded (Fig. 136). Punctation of T2 anteriorly usually not finer than on T1. T3 with coarser and/or more irregular punctation. S2 green, golden or red with often larger black spots (Figs 128–133). Mandible thick or thin	40
39	The following two species are not always separable. Metasoma with slightly convex sides (approximately as in Fig. 100). Head narrower, in frontal view only slightly broader than high. Colour of mesosoma predominantly lighter, often greenish. Usually larger species. Hosts soil-nesting species of Odynerus	C. mediata Linsenmaier
–	Metasoma with more parallel sides (approximately as in Fig. 101). Head narrower, in frontal view distinctly broader than high. Colour of mesosoma predominantly darker, violet, blue or blue-green. Usually smaller species. Hosts cavity-nesting species of Ancistrocerus and Euodynerus	C. solida Haupt
40	Mandible thick (medial width of mandible more than half of its basal width), its margins basally straight in lateral view (Fig. 146). T3 relatively shiny, transgression of lateral margin into lateral apical teeth straight (Fig. 87) or slightly concave (Fig. 88). F1 longer than F2 (Figs 173–174). Metasoma with more parallel sides	41
–	Mandible thinner (medial width of mandible not more than half of its basal width), its margins basally more or less concave in lateral view (Figs 147–149). T3 relatively dull, transgression of lateral margin into lateral apical teeth slightly (Fig. 85) or strongly concave (Fig. 105). F1 longer than or as long as F2 (Figs 175–178). Metasoma with more convex sides	43
41	S2 mainly green, its pubescence dense and long (Fig. 128). F1 distally without or with a very shallow, inconspicuous impression (Fig. 173). Propodeal tooth usually laterally straight or convex	C. corusca Valkeila
–	S2 mainly red or golden, its pubescence variable (Fig. 129). F1 distally with a shallow impression (Fig. 174). Propodeal tooth usually laterally concave	42
42	Pronotum without sharply contrasting border between the darker middle part and the lighter margins (Fig. 160). Punctation of T2 anteriorly coarse or relatively fine. Transgression of lateral margin of T3 into lateral apical teeth straight (Fig. 87). Central interval of apical teeth more angulate, and pits of apical rim smaller (Fig. 87)	C. subcoriacea Linsenmaier
–	Pronotum usually with sharply contrasting border between the darker middle part and the lighter margins (Fig. 161). Punctation of T2 anteriorly coarse. Transgression of lateral margin of T3 into lateral apical teeth slightly concave (Fig. 88). Central interval of apical teeth more arcuate, and pits of apical rim larger (Fig. 88)	C. longula Abeille de Perrin
43	F1 approximately as long as or slightly (not more than 1.2×) longer than F2 (Fig. 175). T2 coarsely punctured throughout (Fig. 105). Punctation of T3 coarse and dense (Fig. 105). Apical teeth of T3 long and sharp (Fig. 105). Pubescense of T3 relatively long. Main colour of sternites green (Fig. 130). Black spots of S2 relatively large and rectangular (Fig. 130). Pubescence of vertex often whitish (Fig. 147)	C. ignita (Linnaeus)
–	F1 1.2–1.5 times as long as F2 (Figs 176–178). T2 more finely punctured, punctures becoming slightly sparser posteriorly (Fig. 107). Punctation of T3 finer (Figs 85, 107). Apical teeth of T3 not as sharp (Figs 85, 107). Pubescense of T3 relatively short. Main colour of sternites variable (Figs 131–133). Black spots of S2 not as rectangular (Figs 131–133). Pubescence of vertex usually brownish (Figs 148, 149)	44
44	The following three species are not always separable. Mandible thinner, its margins basally concave in lateral view (Fig. 148). F1 1.2–1.4 times as long as F3 (Fig. 176). Sternites usually golden green or golden red (Fig. 131). Size of black spots of S2 variable (Fig. 131)	Chrysis schencki Linsenmaier
–	Mandible thicker, its margins basally almost straight in lateral view (Fig. 149). F1 1.2–1.5 times as long as F2 (Figs 177–178)	45
45	F1 1.2–1.4 times as long as F2 (Fig. 177). Size of black spots of S2 variable (Fig. 132). Punctation of mesoscutum coarser, punctures often with lighter colour compared to interstices	Chrysis impressa Schenck
–	F1 1.3–1.5 times as long as F2 (Fig. 178). Black spots of S2 large (Fig. 133). Punctation of mesoscutum finer, punctures not differing in colour from interstices	Chrysis borealis sp. n.

Chrysis varidens-gracillima group

Chrysis gracillima Förster, 1853

Chrysis gracillima Förster, 1853: 328.

Chrysis saussurei Chevrier, 1862: 36.

Diagnosis

Length 4–7 mm. The species is easy to recognise by the edentate posterior margin of T3 and the narrow, elongate body shape. Species of Chrysura, which are similarly coloured and also lack apical teeth, do not have the scapal basin or the dark apical rim, and are larger in size. The head and mesosoma are mainly blue or greenish with golden reflections, and the mesoscutum is medially contrastingly darker than laterally. The metasoma is completely golden red dorsally, but the apical rim is dark blue or blackish. The apical rim is wide, medially slightly undulating and laterally with angled margins. F2, F3 and F4 are ventrally slightly bulging in the male. The shape of the body is very slender and elongate in both sexes.

Distribution

Estonia. Very rare. One female was collected on 14.VII.2015 in Reinu, southwestern Estonia (58.032°N, 24.747°E, leg. V. Soon). No other records are known from the Nordic and Baltic countries, but one female has been collected from Russia, close to the eastern border of Latvia (Pskov Oblast, Krasikovo, 23 km south of Sebezh, VII.1999, leg. A. Reschikov). – West Palearctic: Europe, northern Africa and Middle East (Linsenmaier 1997).

Biology

Habitat: various biotopes with sun-exposed dead wood (Kunz 1994, Linsenmaier 1996, Rosa 2006). Adults often fly on wooden poles (Rosa 2006) and occasionally visit flowers of Apiaceae and Euphorbiaceae (Linsenmaier 1997). Flight period: June to August in Germany (Kunz 1994). Host: species of Microdynerus Thomson (Vespidae) (Friese 1883, Enslin 1929, Benno 1950, Wickl 2001) and possibly also Trypoxylon clavicerum Lepeletier & Serville (Crabronidae) (Morgan 1984). The only Microdynerus species found in Estonia is M. parvulus (Herrich-Schäffer), which could be the host of C. gracillima at the northern limit of its distribution area.

Chrysis succincta group

Chrysis bicolor Lepeletier, 1806

Figs 110, 138

Chrysis bicolor Lepeletier, 1806: 127.

Chrysis succincta var. virideocincta Trautmann, 1927: 160.

Diagnosis

Length 5–8 mm. The species resembles C. illigeri, but the malar space is shorter (Fig. 138), the punctation of T2 is denser and the metascutellum is more raised in profile. Also the black spots of S2 are posteriorly more oblique in the female (Fig. 110). The head and mesosoma are mainly blue or greenish, but the mesoscutum and anterior margin of the pronotum are red or golden red in the female and golden green to greenish in the male. The mesoscutum is usually darkened medially in the male. The metasoma is mainly red in the female and golden greenish in the male with a greenish, bluish or black apical rim. T2 often has a black patch dorsally in the female.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Sweden. Relatively rare. – Trans-Palearctic: Europe, northern Africa, Russian Far East (Kimsey and Bohart 1991, Kurzenko and Lelej 2007).

Biology

Habitat: sparsely vegetated sandy areas. Adults occasionally visit flowers of Apiaceae, Asteraceae, Euphorbiaceae and Rosaceae (Linsenmaier 1997, Rosa 2004, 2006, our own obs.). Flight period: early June to late August. Host: Tachysphex obscuripennis (Schenck) and T. pompiliformis (Panzer) (Crabronidae) (Morgan 1984, Saure 1998, Wickl 2001). In central and southern Europe also Dinetus pictus (Fabricius) (Crabronidae) (Gauss 1967).

Chrysis westerlundi Trautmann, 1927

Fig. 171

Chrysis succincta var. westerlundi Trautmann, 1927: 159.

Chrysis succincta var. nordströmi Trautmann, 1927: 159.

Chrysis westerlundi: Linsenmaier 1959: 113.

Diagnosis

Length 7–9 mm. The species differs from other North European species of the C. succincta group by its characteristic colouration: the head and mesosoma are completely dark blue or almost black dorsally in the female and green blue in the male. The metasoma is dorsally red in both sexes and anteriorly greenish in the male. The metatarsus is long, the second tarsomere is at least 3.5 times as long as broad (Fig. 171). Superficially, the species can resemble similarly coloured species of the C. ignita group, but the two central apical teeth are close to each other and extend further posteriorly than the lateral teeth (as in Fig. 78). Also the black spots of S2 are large and not separated by a metallic central line.

Distribution

Finland. Very rare, only eight specimens (6 females and 2 males) are known. – West Palearctic: the species has been found only from Finland and Russian Fennoscandia (Paukkunen et al. 2014).

Biology

Habitat: sparsely vegetated sandy areas. One specimen was caught with a Malaise trap in a bog. Flight period: mid-June to early August. Host: unknown.

Remarks

The species might be conspecific with the Far Eastern species C. cavaleriei du Buysson, 1908 and/or the North American species C. provancheri Schulz, 1906 (= aurichalcea Provancher, 1881) (Paukkunen et al. 2014).

Figures 165–178.

Metatibia: 165 Chrysis pulcherrima ♀ 166 C. viridula ♀. Mesotibia (arrow indicating spurs): 167 C. pseudobrevitarsis ♀ 168 C. impressa ♀. Hindleg (arrow indicating second tarsomere): 169 C. pseudobrevitarsis ♀ 170 C. impressa ♀ 171 C. westerlundi ♀. Antenna (arrow indicating F1 or F1 and F2): 172 C. brevitarsis ♀ 173 C. corusca ♂ 174 C. subcoriacea ♂ 175 C. ignita ♂ 176 C. schencki ♂ 177 C. impressa ♂ 178 C. borealis sp. n. ♂. Scale 1 mm.

Chrysis illigeri Wesmael, 1839

Figs 78, 111, 139

Habitat: steep sand and loess slopes and stone walls (Ressl 1966, Kusdas 1956, Gerth et al. 2010). Adults occasionally visit flowers of Apiaceae, Caryophyllaceae and Euphorbiaceae (Graffe 1895, Linsenmaier 1997, Rosa 2004). Flight period: June to August. Host: probably Diodontus minutus (Fabricius) and/or Miscophus bicolor Jurine (Crabronidae) (Trautmann and Trautmann 1919, Linsenmaier 1959, Heinrich 1964, Gerth et al. 2010). We consider records of other host species (e.g. Tachysphex nitidus (Spinola)) as uncertain due to lack of supporting information.

Chrysis comparata group

Chrysis scutellaris Fabricius, 1794

Chrysis scutellaris Fabricius, 1794: 458.

Chrysis segmentata Dahlbom, 1829: 9.

Diagnosis

Length 6–9 mm. The species is easily recognised due to its unique colouration among North European chrysidids. The head and the mesosoma are mainly blue or green, but the meso- and metascutellum are red or golden. Usually also the frons, the anterior margin of the pronotum and the lateral sections of the mesoscutum have golden or coppery reflections. The metasoma is dorsally red, but the apical rim of T3 is contrastingly blue. The apical teeth are shallow and indistinct, whereby the apical rim may appear nearly rounded.

Distribution

Denmark, Lithuania, Sweden. Rare. Only a few records from Denmark and Sweden (Scania) and one record from Lithuania are known (Orlovskytė et al. 2010, Sörensson et al. 2012). – West Palearctic: Europe and northern Africa (Linsenmaier 1997, 1999).

Biology

Habitat: xerothermic sparsely vegetated sandy areas, often close to seashore. Adults often bask on sun-exposed pieces of wood or logs on the sand, and spend nights inside hollow plant stems (Sörensson et al. 2012). Occasionally they are found on flowers of Apiaceae, Asteraceae, Crassulaceae and Euphorbiaceae (Trautmann 1927, Heinrich 1964, Linsenmaier 1997, Rosa 2004). Flight period: late June to early August. Host: probably Megachile leachella Curtis (Megachilidae) (Sörensson et al. 2012). In central Europe, also Pseudanthidium lituratum (Panzer) (Megachilidae) (Schmid-Egger et al. 1995).

Remarks

Mitochondrial DNA sequences available at the Barcode of Life Data System (Ratnasingham and Hebert 2007) indicate that Swedish specimens of C. scutellaris differ genetically remarkably from central European specimens. The status of European populations should be studied more in the future.

Chrysis splendidula group

Chrysis splendidula Rossi, 1790

Figs 95, 114

Chrysis splendidula Rossi, 1790: 78.

Diagnosis

Length 5–8 mm. The colouration is similar to C. rutilans, as the head, mesosoma and T3 are green, blue, or partially black, whereas T1 and T2 are golden red. Compared to C. rutilans, the body is more robust and the punctation of T2 is coarser (Fig. 95). T2 has a distinct polished longitudinal keel medially and its posterior margin is narrowly raised (Fig. 95). The black spots of S2 are shorter (Fig. 114) and the head is broader than in C. rutilans.

Distribution

Latvia. Very rare. Only three records are known from eastern and central Latvia. – Trans-Palearctic: Europe, central Asia, Japan and Korea (Linsenmaier 1997, Kurzenko and Lelej 2007).

Biology

Habitat: sparsely vegetated sandy areas, such as sand pits (Hoop 1971). Adults occasionally visit flowers of Apiaceae, Asteraceae and Euphorbiaceae (Linsenmaier 1997, Rosa 2004, 2006). Flight period: July and early August. Host: Eumenes coarctatus (Linnaeus) (Vespidae) (Martynova and Fateryga 2015). Possibly also Eumenes mediterraneus Kriechbaumer and E. pomiformis (Fabricius) (du Buysson 1895, Ferton 1910, Martynova and Fateryga 2015) Other host species reported for C. splendidula, such as Gymnomerus laevipes (Shuckard), Symmorphus allobrogus (Saussure) (Vespidae), Trypoxylon figulus (Linnaeus) (Crabronidae) and Osmia andrenoides Spinola (Megachilidae), concern actually C. rutilans or are unreliable (Martynova and Fateryga 2015).

Chrysis rutilans Olivier, 1791

Fig. 94

Chrysis rutilans Olivier, 1791: 676.

Chrysis splendidula of authors, not Rossi, 1790.

? Chrysis insperata Chevrier, 1870: 265.

Diagnosis

Chrysis nitidula of authors, not Fabricius, 1775.

Chrysis soluta Dahlbom, 1854: 217.

Chrysis purpurata of authors, not Fabricius, 1787.

Diagnosis

Length 7–13 mm. The body is mostly blue or blue-green, resembling C. indigotea in colouration. The female often has green-golden reflections on the mesoscutum, mesoscutellum and anteriorly on the pronotum. The tergites are posteriorly lighter blue than anteriorly. Compared to C. indigotea, the punctation of the mesoscutum is sparser and the interstices larger, the mesoscutum is medially not distinctly darker than laterally, and the ovipositor is narrower (as in Fig. 92). The black spots of S2 are also larger and the apical teeth of T3 shorter and blunter.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Sweden. Relatively rare. – West Palearctic: Europe (Linsenmaier 1997).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Adults are usually found on walls of old log buildings (barns, sheds etc.), log piles, poles and dead tree trunks (e.g. Populus, Salix, Betula). Flight period: late May to late August. Host: Symmorphus allobrogus (Saussure), S. crassicornis (Panzer) and S. murarius (Linnaeus) (Vespidae) (Abeille de Perrin 1878, du Buysson 1895, Pärn et al. 2014, our own obs.).

Chrysis ruddii Shuckard, 1837

Fig. 96

Chrysis Ruddii Shuckard, 1837: 163.

Chrysis auripes Wesmael, 1839: 175.

Diagnosis

Length 7–10 mm. As in most other species of the C. ignita group, the head and mesosoma are mainly blue or green and the metasoma is dorsally golden red. However, the mesoscutum, mesoscutellum and propodeum, and often also tegulae and mesopleuron, have extensive golden or coppery reflections in the female. The sternites and legs are ventrally coppery red in both sexes. The punctation of the tergites is very fine and dense throughout, punctures being of uniform size (Fig. 96). In the male, the punctation is often somewhat sparser, and therefore it is more easily confused with other similarly coloured species of the C. ignita group (e.g. C. subcoriacea). The combination of a short pronotum (length less than one fourth of its width), non-metallic F1 and coppery red sternites should be used to distinguish C. ruddii males reliably from other species of the group.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively common. – West Palearctic: Europe, Asia Minor (Linsenmaier 1997).

Biology

Habitat: dry meadows, rocky outcrops, cliffs, clay banks, forest margins. Adults occasionally visit flowers of Apiaceae and Euphorbiaceae (Heinrich 1964, Linsenmaier 1997, Rosa 2004, 2006). Flight period: mid-May to early August. Host: primarily Ancistrocerus oviventris (Wesmael) (Berland and Bernard 1938, Banaszak 1980, Morgan 1984, Kunz 1994, our own obs.), but possibly also A. parietum (Linnaeus), A. scoticus (Curtis), species of Eumenes Latreille, Odynerus spinipes (Linnaeus) and O. reniformis (Gmelin) (Vespidae) (Forsius in Trautmann 1927, Berland and Bernard 1938, Banaszak 1980, Kunz 1994, Martynova and Fateryga 2015, our own obs.). Records stating solitary bees (e.g. Hoplitis adunca (Panzer) and H. anthocopoides (Schenck)) as hosts are doubtful, as bees differ significantly in their biology from the vespid hosts.

Chrysis corusca Valkeila, 1971

Figs 103, 123, 128, 144, 173

Chrysis corusca Valkeila, 1971: 84.

Diagnosis

Length 7–9 mm. Both sexes are easily confused with e.g. C. angustula, C. schencki, C. impressa and C. longula, and a combination of several characters should always be used in identification. The vertex and mesosoma are mostly dark blue or blue-violet (rarely greenish), and often have green reflections on the pronotum and mesoscutellum. The punctures of the mesoscutum are of the same colour as the interstices. The metasoma is dorsally golden red or violet-red (Fig. 103) and S2 is greenish/bluish with narrow black spots (Figs 123, 128). The shape of the body is elongate and nearly parallel-sided (Fig. 103). Punctation of T2 is relatively regular and coarse (Fig. 103) and the mandible is thick (Fig. 144). The apical rim of T3 has a characteristic, widely arcuate interval between the central apical teeth.

Distribution

Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – West Palearctic: central and northern Europe, Iran (van der Smissen 2010, Rosa et al. 2013).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Rarely also sandy banks and clay structures, such as old barn walls. Flight period: early June to early August. Host: Symmorphus gracilis (Brullé) (Vespidae) (Pärn et al. 2014).

Chrysis clarinicollis Linsenmaier, 1951

Figs 98, 162

Chrysis ignita var. clarinicollis Linsenmaier, 1951: 77.

Chrysis clarinicollis: Schmid-Egger et al. 1995: 267.

Diagnosis

Length 6–10 mm. The species is characterised by a uniformly green or blue-green pronotum and mesoscutellum, and a contrastingly darker blue or violet mesoscutum (Fig. 162). The tergites are mainly golden red, but T1 is anteriorly and laterally blue-green or green (Fig. 98). Shape of the metasoma is broad and compact (Fig. 98). Punctation of T2 is somewhat finer than on T1 and dense throughout (Fig. 98). The sternites and legs are ventrally greenish. The apical rim of T3 is narrow and the intervals of the apical teeth very shallow (Fig. 98). The mandible and ovipositor are thin (as in Figs 92 and 141). Males in particular can be confused with other species of the C. ignita group as their colouration is more variable than in females. The colouration should always be used in combination with other characters (e.g. shape of metasoma and apical teeth) in species identification.

Distribution

Estonia. Very rare. Only four records are known from Estonia, all from the western part of the country (Ahu, Manilaid, Tuudi, Väike-Pakri; 2001–2009). – West Palearctic: southern and central Europe, northern Africa (Linsenmaier 1997).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Usually observed on dead tree trunks and on walls of abandoned houses (Rosa 2006), but also flying near the ground and over rocks (Linsenmaier 1997). Flight period: June to August. The earliest Estonian specimens were collected in the beginning of June and the latest in the end of July. Host: possibly Ancistrocerus oviventris (Wesmael) and/or A. scoticus (Curtis) (Vespidae) (Petit 1987).

Chrysis vanlithi Linsenmaier, 1959

Figs 97, 115, 140, 159

Chrysis rutiliventris ssp. vanlithi Linsenmaier, 1959: 153.

Chrysis rutiliventris of authors, not Abeille de Perrin, 1879.

Chrysis vanlithi: Soon et al. 2014: 305.

Diagnosis

Length 7–10 mm. The species is easily confused with other similarly coloured species of the C. ignita group (e.g. C. borealis sp. n. and C. schencki), and the males in particular can be difficult to identify. The combination of several characters (e.g. shape of pronotum and malar space and colouration) should always be used in species determination. The head and mesosoma are dorsally dark blue or nearly black with light blue or greenish reflections mainly on the pronotum (Fig. 159). The tergites are golden red (Fig. 97) and the sternites and legs ventrally greenish (Fig. 115). The pronotum is short (length not more than one fourth of width) (Fig. 159). The mandible is relatively thin (Fig. 140) and the malar space long, approximately as long as broad in the female (Fig. 140). F1 is black without a metallic sheen, and the ovipositor is narrow (as in Fig. 92).

Distribution

Denmark, Norway, Sweden. Rare. Only one confirmed record is known from Denmark, four from southern Sweden (Scania, Bohuslän and Stockholm archipelago) and ten from southern Norway. – West Palearctic: from central and northern Europe to southwestern Asia (Linsenmaier 1997).

Biology

Habitat: all Nordic specimens have been found in coastal localities. Adults are usually found flying near rocks and log walls (Linsenmaier 1997, our own obs.). Flight period: June to July (most Nordic observations are from June). Host: possibly Ancistrocerus gazella (Panzer) (our own obs.) or Ancistrocerus scoticus (Curtis) (Vespidae) (van der Smissen 2001).

Chrysis subcoriacea Linsenmaier, 1959

Figs 87, 109, 117, 129, 146, 160, 174

Chrysis longula ssp. subcoriacea Linsenmaier, 1959: 160.

Chrysis subcoriacea: Valkeila 1962: 64.

Diagnosis

Length 9–13 mm. Females are usually easy to recognise by the laterally coriaceous and dull T2 and T3 (Fig. 109). Males, however, are often confused with C. impressa and C. longula. Compared to C. impressa, the sternites are usually brighter red and the black spots of S2 larger (Fig. 129), the mandible thicker (Fig. 146) and the transition of the lateral margin of T3 into the lateral apical teeth is straighter (not concave) (Fig. 87). Compared to C. longula (Fig. 161), the middle part of the pronotum is not as sharply darker than the margins (Fig. 160), the lateral margins of T3 are straighter, the punctation of T2 is usually finer anteriorly, and the central interval of the apical teeth is more angled (Fig. 87). In both sexes, the head and mesosoma are dorsally dark blue or nearly black with light blue or greenish reflections mainly on the pronotum. The tergites and sternites are red. The body is elongate and often rather large.

Distribution

Denmark, Estonia, Finland, Latvia, Norway, Sweden. Relatively rare. – Trans-Palearctic: from Europe to central Asia and Japan (Linsenmaier 1997).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Adults often fly near dead tree trunks (Betula, Populus, Salix) and close to walls of wooden buildings (log barns, sheds etc.), and they are also attracted to honeydew of aphids. Flight period: late May to late August. Host: possibly Ancistrocerus trifasciatus (Müller) (Vespidae) (our own obs.).

Chrysis angustula Schenck, 1856

Figs 86, 99, 119, 126

Chrysis angustula Schenck, 1856: 28.

? Chrysis gracilis Schenck, 1856: 30.

Chrysis brevidens Tournier, 1879: 96.

Diagnosis

Length 6–9 mm. The species can be confused with several other similarly coloured species of the C. ignita group (e.g. C. leptomandibularis, C. schencki and C. corusca). Compared to C. leptomandibularis and C. schencki, the mandible is thicker, the punctation of T2 is finer, the black spots of S2 are more rectangular and the posterior margin of the propodeal tooth is directed more downward. Compared to C. corusca, the mandible is thinner, the punctation of the tergites is finer and the colour of the sternites is more reddish (not green). The head and mesosoma are mainly dark blue, and usually have extensive green or golden reflections on the frons, pronotum, mesopleuron, mesoscutum and mesoscutellum, especially in the female. The metasomal tergites are golden red (Fig. 99) and the sternites golden or reddish (Figs 119, 126). The black spots of S2 are characteristically rectangular in shape (Figs 119, 126). The body is elongate and slender, with parallel sides (resembling C. leptomandibularis) (Fig. 99). The punctation of T2 is fine, and its surface is strongly shining posteriorly (Fig. 99). T3 is relatively long, and strongly shining, especially in the female (Fig. 99). The apical teeth are short, and the central interval is wide and shallow (Figs 86, 99). The ovipositor is narrow (as in Fig. 92) and the mandible relatively thick in both sexes.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Very common. – Trans-Palearctic. Europe, southwestern Asia, Siberia, China (Linsenmaier 1997, Rosa et al. 2014).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Adults are often found on walls of wooden buildings, poles, log piles and dead tree trunks. Occasionally they also visit flowers of Apiaceae and Asteraceae (Banaszak and Kochanowski 1994, our own obs.) and are attracted to honeydew of aphids. Flight period: late May to September. Host: primarily Symmorphus bifasciatus (Linnaeus) (van Lith 1958, Niehuis 2000, Pärn et al. 2014, our own obs.), but occasionally also Ancistrocerus trifasciatus (Müller), S. allobrogus (Saussure), S. connexus (Curtis) and S. debilitatus (Saussure) (Vespidae) (van Lith 1958, Niehuis 2000, Pärn et al. 2014, our own obs.).

Chrysis longula Abeille de Perrin, 1879

Figs 6, 88, 102, 121, 161

Chrysis ignita var. longula Abeille de Perrin, 1879: 74.

Chrysis longula var. sublongula Linsenmaier, 1951: 76.

Chrysis longula ssp. aeneopaca Linsenmaier, 1959: 160.

Chrysis longula: Linsenmaier 1959: 159.

Diagnosis

Length 10–13 mm. The body is elongate with parallel sides (Fig. 102) and usually large compared to other species of the C. ignita group. The head and mesosoma are dorsally blue or black, and the female has extensive golden green reflections on the pronotum, mesopleuron and mesoscutellum. The punctures of the mesoscutum are usually lighter coloured than the interstices (as in C. impressa). The tergites and sternites are golden red (Figs 102, 121) and the black spots of S2 are long and narrow (Fig. 121). The punctation of T2 is anteriorly very coarse, and the surface of T3 is shiny in the female (Fig. 102). The mandible is long and relatively thick (in the male as in Fig. 146). Small specimens can be confused with C. angustula and C. corusca, but the punctation of T2 is coarser anteriorly and the black spots of S2 are narrower. The sternites are mostly red, not greenish as in C. corusca. The males are often difficult to distinguish from C. impressa and C. subcoriacea. Compared to C. impressa, the sternites are usually brighter red and the black spots of S2 are more elongate. The punctation of T2 is coarser anteriorly and the metasoma is more elongate in shape. Compared to C. subcoriacea, the pronotum is more abruptly bicoloured (Fig. 161), the punctation of T2 is usually coarser anteriorly, the lateral margin of T3 is more concave and the central interval of the apical teeth is more arcuate (Fig. 88).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – Trans-Palearctic: from western Europe to central Asia, Siberia and China (Linsenmaier 1959, 1997, Rosa et al. 2014).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Adults can be found on walls of old log buildings (barns, sheds etc.), log piles, poles and dead tree trunks (e.g. Betula, Populus, Salix). Flight period: early June to late August. Host: Ancistrocerus antilope (Panzer), Symmorphus crassicornis (Panzer) and S. murarius (Linnaeus) (Linsenmaier 1959, Heinrich 1964, Morgan 1984, Brechtel 1985, Petit 1987, Martynova and Fateryga 2015), possibly also Ancistrocerus parietinus (Linnaeus) and species of Euodynerus Dalla Torre (Vespidae) (Blüthgen 1961, Morgan 1984, Martynova and Fateryga 2015).

Remarks

Eastern Palearctic populations of C. longula belong to ssp. aeneopaca Linsenmaier, 1959 which differs from the nominotypical subspecies by having fine punctation and brownish colour anteriorly on tergites. Specimens of ssp. aeneopaca can sometimes be confused with C. subcoriacea. One specimen similar to ssp. aeneopaca has been found in Finland (Kuopio), but the occurrence of the taxon in Fennoscandia is questionable (Paukkunen et al. 2014).

Chrysis brevitarsis Thomson, 1870

Figs 155, 157, 172

Estonia, Latvia, Lithuania, Sweden. Relatively rare. – West Palearctic? Reliable distributional data from the eastern Palaearctic are not available. Records from Japan have been found to be erroneous (Linsenmaier 1997).

Biology

Habitat: sun-exposed clay and loess walls and banks, sand pits, road verges and meadows. Adults have been found on flowers of Euphorbiaceae (Rosa 2004). Flight period: early June to late July. Host: primarily Odynerus spinipes (Linnaeus), but also O. reniformis (Gmelin) (Vespidae) (van Lith 1958, Linsenmaier 1959, Banaszak 1980, Morgan 1984).

Chrysis solida Haupt, 1957

Figs 91, 101, 120, 125, 137

Chrysis ignita ssp. solida Haupt, 1957: 115.

Chrysis mediata ssp. fenniensis Linsenmaier, 1959: 154.

Chrysis mediata of authors, not Linsenmaier, 1951.

Chrysis scintillans Valkeila, 1971: 85.

Chrysis solida: Niehuis 2001: 120.

Diagnosis

Remarks

Recent mitochondrial DNA studies have shown that C. schencki consists of two distinct and sympatric genetic lineages in northern Europe (Soon et al. 2014). It is possible that they represent two different species, but more morphological and molecular studies are needed.

Chrysis ignita (Linnaeus, 1758)

Figs 104, 105, 122, 130, 147, 163, 175

Sphex ignita Linnaeus, 1758: 571.

Chrysis ignita: Linnaeus 1761: 414.

Chrysis ignita form B sensu Linsenmaier 1959: 156.

Diagnosis

Length 5–10 mm. C. ignita resembles closely C. terminata in colouration, structure and habitus, but the frontal carina is shallowly M-shaped or more or less arcuate (not forming four tooth-like tubercles). The head and mesosoma are dorsally shiny blue or violet with green reflections on the pronotum and mesoscutellum (Fig. 163). The punctures of the mesoscutum are of the same colour as the interstices (Fig. 163) (not lighter as in C. impressa). The tergites are golden red (Figs 104, 105), the sternites green or blue (Figs 122, 130) and the black spots of S2 are subrectangular in shape (Figs 120, 130). The punctation of T2 and T3 is coarse and regular (Figs 104, 105) and the apical teeth are sharply produced (Figs 104, 105). The medial furrow of the pronotum is narrow (Fig. 163) and the pubescence of the vertex is white (Fig. 147) or sometimes brown in the male.

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Relatively rare. – West Palearctic: from western Europe to central Asia and China (Linsenmaier 1997, Rosa et al. 2014).

Biology

Habitat: gardens, parks and forest margins. Adults are usually collected from walls of old buildings (both wooden and stone), dead tree trunks, poles and log piles. They rarely visit flowers of Apiaceae (Rosa 2004). Flight period: late May to early September. Host: probably Ancistrocerus parietum (Linnaeus) (Vespidae) (our own obs.). Numerous host records have been published for C. ignita, but most of these are unreliable, due to inconsistent taxonomic treatment of the species.

Remarks

A few studied specimens from Norway, Finland and Lithuania differ significantly from other North European C. ignita specimens based on their mitochondrial DNA sequences. According to Soon et al. (2014), they could represent a cryptic species (“Chrysis sp.1”). No distinct morphological differences have been found between the two North European genetic forms.

Chrysis impressa Schenck, 1856

Figs 10, 84, 85, 92, 107, 108, 116, 132, 149, 164, 168, 170, 177

Chrysis impressa Schenck, 1856: 29.

Chrysis ignita var. aurifera Linsenmaier, 1951: 76.

Diagnosis

Length 6–11 mm. The species is easily confused with other similarly coloured species of the C. ignita group and a combination of different diagnostic characters should be used in species determination. The head and the mesosoma are dorsally dark blue or black, and in the female the pronotum, mesopleuron and mesoscutellum have extensive golden green reflections (Fig. 164). The mesoscutum of the female is characteristically black, dark grey or olive coloured with contrastingly green or blue punctures (similar to C. longula) (Fig. 164). The mesoscutum of the male is often entirely dark blue or blue-violet. The tergites are golden red and relatively finely punctured (Figs 107, 108). The sternites are at least partially red-golden (Figs 116, 132) and the black spots of S2 are usually roundish in the female (Figs 116). The setae on the dorsal surface of the head are brownish in both sexes (Fig. 149). The mandible is relatively thick (medial width about half or nearly half of basal width) and basally with only slightly concave margins (Figs 141, 149). F1 is long and narrow, about 1.4 times as long as F2 in the female and at least 1.2 times as long as F2 in the male (Fig. 177).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Very common. – West Palearctic: from western Europe to central Asia (Linsenmaier 1997).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Adults are mainly observed flying and running on walls of wooden buildings (e.g. log barns), dead tree trunks (e.g. Betula, Populus), poles and log piles. Flight period: early June to late August. Host: Mainly Ancistrocerus claripennis Thomson and A. parietinus (Linnaeus) (Pärn et al. 2014, Martynova and Fateryga 2015, our own obs.), but probably also A. trifasciatus (Müller) (Vespidae) (Morgan 1984, Pärn et al. 2014, our own obs.).

Chrysis borealis Paukkunen, Ødegaard & Soon, sp. n.

http://zoobank.org/3DDF3EF4-6694-4586-B78E-FF79E321AB92

Figs 106, 124, 133, 136, 158, 178, 179, 180–186, 187–195

Chrysis mediadentata of authors, not Linsenmaier, 1951.

Chrysis sp. sensu Soon et al. (2014) and Paukkunen et al. (2014: 44).

Type material

11 ♀♀ and 15 ♂♂. DNA barcode sequences of all type specimens are available at GenBank or Barcode of Life Data System (Ratnasingham and Hebert 2007).

Holotype

Norway, ♀ (HYMNI560), Nord-Trøndelag, Ørin, 63.802°N, 11.459°E, 9.VII.2014, leg. F. Ødegaard (NUM) (Fig. 179).

Paratypes

Finland, 1 ♂ (FACU-000396), Kittilän Lappi, Kolari, Ylläs, 67.586°N, 24.239°E, 9.VII.1989, leg. M. Koponen (MZH); 1 ♂ (MZH_GP.74723), Sompion Lappi, Sodankylä, 68.027°N, 27.413°E, 1.VII.2002, leg. J. Itämies (MZH); 1 ♀ (TUZ616001), Enontekiön Lappi, Malla Strict Nature Reserve, 69.060°N, 20.759°E, 15.VII.2009, leg. V. Soon (TUZ); 1 ♀ (FACU-000399), Enontekiön Lappi, Annjaloanji, 69.172°N, 21.439°E, 11.VII.2007, leg. R. Jussila (private collection of M. Raekunnas, Hämeenlinna, Finland); 2 ♂♂ (MZH_GP.92704, MZH_GP.92705), Inarin Lappi, Inari, Ivalo, 68.643°N, 27.524°E, 26.VI.2013, leg. T. Järveläinen (MZH and NMLS); Norway, 1 ♂ (Chrysis132), Buskerud, Hokksund, Lilleby, 59.779°, N 9.933°E, 1.V.2012, leg. F. Ødegaard (NUM); 1 ♂ (Chrysis004), Buskerud, Nedre Eiker, Solbergfjell, 59.759°N, 10.041°E, 28.VI.2012, leg. F. Ødegaard (NUM); 1 ♀ (NOCHR254), Hordaland, Masfjorden, Mjanger, 60.768°N, 5.348°E, 3.IX.2009, leg. A. Staverløkk (NUM); 1 ♂ (HYMNI559), Nord-Trøndelag, Ørin, 63.802°N, 11.459°E, 9.VII.2014, leg. F. Ødegaard (NUM); 1 ♂ (Chrysis034), Oppland, Nord-Fron, Stordalsberget, 61.587°N, 9.819°E, 9.V.2009, leg. F. Ødegaard (NUM); 1 ♂ (Chrysis089), Oppland, Nord-Fron, Stordalsberget, 61.587°N, 9.819°E, 1.VI.2009, leg. F. Ødegaard (NUM); 1 ♀ (Chrysis180), Oppland, Nord-Fron, Stordalsberget, 61.587°N, 9.819°E, 1.IX.2009, leg. F. Ødegaard (NUM); 1 ♂ (Chrysis183), Sogn og Fjordane, Luster, Ornes, 61.286°N, 7.341°E, 4.VII.2011, leg. F. Ødegaard (NUM); 1 ♂ (Chrysis021), Sør-Trøndelag, Røros, Småsetran, 62.573°N, 11.413°E, 11.VII.2010, leg. F. Ødegaard (NUM); 1 ♀ (Chrysis187), 1 ♂ (Chrysis189), Sør-Trøndelag, Røros, Småsetran, 62.573°N, 11.413°E, 23.VII.2007, leg. F. Ødegaard (NUM); 1 ♂ (Chrysis125), Sør-Trøndelag, Røros, Kvitsanden, 62.573°N, 11.412°E, 31.VII.2008, leg. F. Ødegaard (NUM); 1 ♂ (NOCHR267), Sør-Trøndelag, Trondheim, Lade, 63.447°N, 10.434°E, 27.V.2013, leg. F. Ødegaard (NUM); Russia, 1 ♀ (MZH_GP.78002), Lapponia tulomensis, 45 km east of Murmansk, 68.876°N, 34.196°E, 16.VII.2006, leg. M.V. Kozlov (MZH); Sweden, 1 ♀ (TUZ616002), Öland, Persnäs, 57.046°N, 16.931°E, 20.VII.2007, leg. J. Abenius (TUZ); 1 ♀ (MZH_GP.92690), Gotland, Stora Karlsö, Hien, 57.289°N, 17.964°E, 6.VII.2012, leg. N. Johansson (MZH); 1 ♀ (MZH_GP.92688), 1 ♂ (MZH_GP.92689), Gotland, Stora Karlsö, Hien, 57.289°N, 17.964°E, 8.VII.2012, leg. N. Johansson (MZH); 1 ♀ (MZH_GP.92691), Gotland, Fårö, Norsta Auren, 57.981°N, 19.326°E, 8.VII.2012, leg. N. Johansson (MZH).

Diagnosis

Length 6–11 mm. The species is very similar to C. impressa in shape and structure, but the colouration is darker and the length of F1 compared to F2 is larger (Table 1). The mesoscutum of the female is usually black, violet or dark blue with relatively fine and dense punctation (Figs 158, 179). The punctures are generally of the same colour as the interstices. The metasoma has golden red or reddish tergites (Figs 106, 179) and the sternites are dark green or bluish in the female (Fig. 124), but often with golden red colour in male (Fig. 133). Compared to C. corusca, the body shape is stouter, the metasoma is notably swollen (Figs 106, 179), the black spots of S2 are broader (Figs 124, 133) and the mandible is thinner. Dark specimens of C. schencki are also very similar, but have a thinner mandible and coarser punctation on the scapal basin, and are more slender in habitus. The males of C. borealis in particular are difficult to distinguish from C. impressa and C. schencki. On average, the length ratio F1/F2 is larger (1.3–1.5:1) (Fig. 178, Table 1), the black spots of S2 are larger (Fig. 133) and the punctation of the mesoscutum is finer in C. borealis. Identification of the males is not always possible with certainty.

Description of female

Body length 7.8–10.3 mm, forewing length 5.1–6.6 mm (n = 12).

Head. Height 1.8–2.1 mm, width 2.3–2.6 mm, length 1.0–1.1 mm, shortest interocular distance 9.4–11.0 mm. Scapal basin green or greenish blue, usually becoming darker blue or violet dorsally below frontal carina. Punctation of scapal basin very dense and fine, partially coriaceous with rugae formed by the puncture margins. Transverse frontal carina well developed, usually relatively evenly arcuate or slightly notched medially. Vertex dark blue, dark violet or black. Pubescence on vertex light brownish. Malar space 1.4 times broader than high. Mandible blackish brown, apically and in inner margin light brown, without subapical tooth. In lateral view, mandible relatively thick (similar to C. impressa), its sides medially almost parallel and basally only slightly concave. Scapus, pedicellus, and F1 with green, blue or violet metallic reflections. Relative lengths of P/F1/F2/F3 are 1/1.8/1.1/0.9. F1 usually 1.4–1.7 times as long as F2 (Table 1). F4–F10 approximately 1.2 times as long as broad.

Mesosoma. Length 3.0–3.8 mm, width anterior to tegulae 2.0–2.7 mm. Length of pronotum medially 0.5–0.6 mm and width at anterior margin 1.8–2.2 mm. Colour of pronotum medially black, dark violet or dark blue, on the margins lighter green, blue or violet, only rarely with golden reflections (Figs 158, 179). Medial groove relatively shallow and indistinctly delimited. Mesoscutum dark violet or black, sometimes with bluish reflections (Figs 158, 179). Punctures generally of the same colour as interstices. Punctation of mesoscutum relatively fine and dense with narrow interstices (Figs 158, 179). Size of punctures on average smaller than in C. impressa and C. schencki. Interstices with scattered small punctures. Tegula green, blue or violet, with paler colour laterally. Mesoscutellum black medially and violet, blue, or sometimes greenish laterally, with irregular large punctures. Metanotum and propodeum violet, blue or greenish. Outer margin of lateral propodeal teeth straight or slightly concave. Pubescence on mesosoma whitish. Legs violet, blue or greenish, but tarsal segments dark brown. Wing venation as in C. impressa and C. schencki.

Metasoma. Length 3.8–4.9 mm, maximum width 2.4–2.9 mm. Colour of tergites golden red or reddish, T1 anteriorly often greenish (Fig. 106). T1 with strong punctation, a weakly elevated medial line with sparser punctation, and very small and scattered punctures on interstices. Punctation of T2 anteriorly regular and dense, of the same strength as on T1 (Fig. 106). Punctation becoming weaker and more scattered laterally and posteriorly. T2 with prominent elevated medial line, anteriorly narrow and posteriorly broad, flat and shiny. T3 weakly saddle-shaped, with regular, strong and dense punctation, and often with elevated midline. Punctures of T3 on average as large as posteriorly on T2. Interstices usually with prominent microsculpture, whereby surface appears dull (Fig. 106). Posterior margin of T3 with four broadly separated teeth, intervals shallow. Medial interval usually about 1.5 times wider than lateral intervals. Subapical pit row with 15–20 black or bluish pits. Subapical lateral swellings relatively strong. Pubescence silvery white. Sternite colour green or blue, occasionally with golden reflections (Fig. 124). Black spots of S2 relatively large and subrectangular, their margins often vaguely delimited (Fig. 124). Ovipositor thin, similar to C. ignita and C. impressa (Fig. 92). Internal sternites and tergites similar to C. impressa (Figs 180–186). T5 relatively narrow, about three times as long as broad, and tapering posteriorly, without lateral stigmae (Fig. 181).

Description of male

Body length 6.7–9.0 mm, forewing length 4.7–6.0 mm (n = 15).

Head. Height 1.5–2.0 mm, width 1.9–2.4 mm, length 0.8–1.1 mm, shortest interocular distance 0.3–0.4 mm. Structure and colouration as in female, but scapal basin often slightly paler, shape of transverse frontal carina more variable, pubescence longer and mandible thicker. Sides of mandible basally slightly concave, gradually converging towards apex in lateral view. Relative lengths of P/F1/F2/F3 are 1/1.8/1.3/1.2 (Fig. 178). F1 usually 1.3–1.5 times as long as F2 (Table 1). F4–F10 as in female, or slightly shorter.

Mesosoma. Length 2.5–3.5 mm, width anterior to tegulae 1.7–2.4 mm. Length of pronotum medially 0.3–0.6 mm and width at anterior margin 1.5–2.1 mm. Structure as in female, but colouration usually somewhat lighter and pubescence longer. Margins of pronotum more often with golden reflections, and mesoscutum sometimes entirely blue. Mesoscutellum often medially violet, not always black, whereas mesoscutellum laterally, metanotum and propodeum violet, blue or golden green. Legs green, golden green or bluish with dark brown tibiae.

Metasoma. Length 3.3–4.4 mm, maximum width 2.1–2.8 mm. Colour of tergites as in female, but punctation of T1 and T2 usually denser and finer. T3 with very dense and homogenous punctation. Interstices shining without distinct microsculpture. T3 convex, not medially depressed as in female. Shape of apical teeth of T3 relatively variable. Medial interval narrower than or as wide as lateral intervals. Subapical pit row with 12–20 black pits. Subapical lateral swellings weak or nearly missing. Sternites with green, golden and reddish colour (Fig. 133), sometimes almost completely green. Black spots of S2 large and rounded (Fig. 133). Inner margin of paramere rounded (Fig. 136). Internal sternites and tergites similar to C. impressa (Figs 187–195). S8 about 1.2 times as long as broad, posteriorly pointed and anteriorly rounded (Fig. 195).

Geographic variation

Southern specimens from Estonia, Öland and Gotland are more uniform in colour than specimens from Finland, Norway and the Swedish mainland. The mesosoma of southern specimens is uniformly bright blue or violet with some greenish reflections, whereas in northern specimens, the mesoscutum and central part of the pronotum are commonly black or dark violet, and the margins of the pronotum and mesoscutellum are, in contrast, greenish or even golden green, especially in the males.

DNA analysis

Variable positions of the DNA barcode sequences of C. borealis sp. n. and its sibling species, C. ignita and C. impressa, are presented in Table 2. Despite relatively high intraspecific variability, there are two diagnostic nucleotide mutations conserved in all sequences of C. borealis sp. n. compared to C. ignita and C. impressa: G instead of A in position 241 and T instead of C in position 340. Additionally, there are three transitions shared with C. impressa, but differing from C. ignita, and one transition shared with C. ignita, but differing from both haplotypes of C. impressa. All samples of C. borealis sp. n. cluster together indicating their closer relationship with each-other than the other two species (Fig. 196).

Distribution

Denmark, Estonia, Finland, Norway, Sweden. Rare. – West Palearctic (?), general distribution poorly known. So far only known from the Nordic and Baltic countries and north-western Russia (Leningrad Oblast, Republic of Karelia, Murmansk Oblast) (Paukkunen et al. 2014).

Biology

Habitat: rocky outcrops, cliffs, alpine meadows, forest margins. Often found on islands of the Baltic Sea and in Lapland, where other species of the C. ignita group are uncommon. Adults have been found sitting on sun-exposed leaves of Tussilago and flowers of Apiaceae. They have also been collected using yellow pan traps. Flight period: late May to late August. A few specimens have been collected in early May and September. Host: Ancistrocerus parietum (Linnaeus) (Vespidae), based on records of C. borealis sp. n. from islands and other coastal localities, where A. parietum is the only species of Eumeninae. In northern alpine areas, where A. parietum is not present, A. scoticus (Curtis) is the most likely host species.

Etymology

The species epithet borealis is a Latin word derived from the Greek boreas which means north. We use it here as an adjective in the feminine case. The interpretation of borealis should be “northern”.

Remarks

C. borealis sp. n. is very closely related to C. impressa, and cannot always be determined with certainty by morphological characters only. It is also easily confused with C. schencki. The colouration of C. impressa can sometimes be relatively dark and similar to C. borealis n. sp., which possibly could be caused by cool weather during the larval and pupal development. Generally, the colouration of chrysidids becomes darker in northern and alpine localities with cool climatic conditions. C. borealis n. sp. is mainly found in cooler habitats than C. impressa, so it could be claimed to constitute only a dark ecological form of C. impressa and not a distinct species. The slight, but constant divergence in the DNA barcode sequence (Table 2) and the statistically significant difference in the F1/F2 ratio (Table 1), however, supports the treatment of C. impressa and C. borealis n. sp. as distinct, but evolutionarily young sibling species. It is also noteworthy that the DNA barcode sequence of C. ignita is very similar to C. borealis n. sp. and C. impressa (Table 2), but due to significant morphological and ecological differences C. ignita undoubtedly forms a distinct species. Previous studies indicate that morphological and molecular differences between biologically well-defined species can be extremely small or even nonexistent in the C. ignita group (Soon et al. 2014). An example is provided by C. mediata and C. solida, which are not always reliably separable by morphological characters or DNA barcodes, but represent ecologically well separated species with different hosts and habitats (Soon et al. 2014).

Several authors have earlier identified specimens of C. borealis sp. n. erroneously as C. mediadentata Linsenmaier, 1951 (Paukkunen et al. 2014). For example, Erlandsson (1971) reported C. mediadentata from Sweden, Norway and eastern Fennoscandia based on such misidentified specimens. The dark colouration of C. mediadentata can indeed resemble that of C. borealis sp. n., but the species is easily distinguished by the breadth of the ovipositor, which in C. borealis sp. n. is of the thin ignita-type (as in Fig. 92), and in C. mediadentata of the thick solida-type (as in Fig. 91) (Linsenmaier 1987). Additionally, the apical teeth of C. mediadentata have deeper intervals, and the two central teeth are more narrowly separated. The mandible of C. mediadentata is thicker and the inner margin of the paramere of the male genitalia is angulate (as in Fig. 137). So far C. mediadentata has not been found in the Nordic and Baltic countries. The northernmost confirmed records are from northern Germany (Jacobs and Kornmilch 2007).

Figure 179.

Holotype of Chrysis borealis sp. n. ♀ (HYMNI560) Collected from Ørin, Nord-Trøndelag, Norway. Scale 1 mm. (Photo: Arnstein Staverløkk).

Table 1.

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Length ratio of the first and second flagellomere in Chrysis borealis sp. n. and closely related species. All specimens of C. borealis sp. n. and C. impressa, and most of C. schencki were identified by DNA barcoding. In C. ignita, most specimens were identified by morphological characters only. A T-test was used for studying statistical differences between species.

Species	Sex	N	F1/F2				P-value of T-test
Species	Sex	N	Mean	Sd	Max	Min	C. borealis sp. n.	*C. ignita*	*C. impressa*
C. borealis sp. n.	male	16	1.38	0.07	1.52	1.25	-	-	-
C. ignita	male	10	1.17	0.05	1.25	1.12	<0.0001	-	-
C. impressa	male	18	1.27	0.07	1.41	1.14	<0.0001	0.0001	-
C. schencki	male	10	1.26	0.09	1.39	1.11	0.0008	0.0078	0.3314
C. borealis sp. n.	female	9	1.58	0.11	1.73	1.42	-	-	-
C. ignita	female	10	1.41	0.07	1.50	1.29	0.0005	-	-
C. impressa	female	13	1.44	0.11	1.63	1.25	0.0045	0.1582	-
C. schencki	female	10	1.48	0.07	1.58	1.38	0.0128	0.0165	0.1916

Figures 180–186.

Internal tergites and sternites of Chrysis borealis sp. n. ♀: 180 T4 181 T5 182 T6 183 T7 184 S4 185 S5 186 S6. Illustrations based on a paratype from Sweden (MZH_GP.92691). Scale 1 mm. (Drawings: Juho Paukkunen)

Figures 187–195.

Internal tergites and sternites of Chrysis borealis sp. n. ♂: 187 T4 188 T5 189 T6 190 T7 191 S4 192 S5 193 S6 194 S7 195 S8. Illustrations based on a paratype from Finland (MZH_GP.92704). Scale 1 mm. (Drawings: Juho Paukkunen)

Table 2.

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Variable positions in DNA barcode sequences of Chrysis borealis sp. n. (type specimens) compared with all known haplotypes of C. ignita and C. impressa (sensu Soon et al. 2014). Dots (.) indicate similarity with the reference sequence on the first line, hyphens (-) indicate missing data. Numbering corresponds to the COI barcode standard used for insects.

					1	1	2	2	2	3	3	3	3	3	4	6	6	6
	4	5	5	7	3	8	1	1	4	3	4	4	6	6	6	2	3	4
	3	0	5	7	3	4	2	4	1	7	0	3	1	4	9	8	4	6
C. borealis sp. n. NOR (Chrysis183)	G	T	G	G	A	G	C	A	G	T	T	A	C	T	T	A	T	T
C. borealis sp. n. NOR (HYMNI560)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	-	-	-
C. borealis sp. n. NOR (NOCHR267)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	-
C. borealis sp. n. NOR (Chrysis034)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	-
C. borealis sp. n. NOR (Chrysis021)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. NOR (Chrysis089)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. NOR (Chrysis187)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. SWE (MZH_GP.92690)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. SWE (MZH_GP.92688)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. SWE (MZH_GP.92689)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. FIN (MZH_GP.92704)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. RUS (MZH_GP.78002)	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.	.
C. borealis sp. n. FIN (MZH_GP.92705)	.	.	.	.	.	.	.	.	.	.	.	.	.	C	.	.	.	C
C. borealis sp. n. NOR (Chrysis125)	.	.	.	.	.	.	.	T	.	.	.	.	.	.	.	.	.	C
C. borealis sp. n. FIN (MZH_GP.74723)	.	.	.	.	.	.	.	T	.	.	.	.	.	.	.	.	.	C
C. borealis sp. n. FIN (FACU-000396)	.	.	.	.	.	.	.	T	.	.	.	.	.	.	.	.	.	C
C. borealis sp. n. SWE (TUZ616002)	.	C	.	.	.	.	.	.	.	.	.	.	T	.	.	.	.	.
C. borealis sp. n. NOR (NOCHR254)	.	.	.	.	.	.	.	.	.	.	.	.	T	.	.	.	.	-
C. borealis sp. n. NOR (Chrysis180)	.	.	.	.	.	.	.	.	.	.	.	.	T	.	.	.	.	.
C. borealis sp. n. NOR (Chrysis132)	.	.	A	.	.	.	.	T	.	.	.	.	T	.	.	.	.	.
C. borealis sp. n. NOR (Chrysis004)	.	.	A	.	.	.	.	T	.	.	.	.	T	.	.	.	.	.
C. borealis sp. n. SWE (MZH_GP.92691)	.	.	A	.	.	.	.	T	.	C	.	.	T	.	.	.	.	.
C. borealis sp. n. FIN (TUZ616001)	.	.	A	.	.	.	.	.	.	.	.	.	T	.	C	.	.	.
C. borealis sp. n. FIN (FACU-000399)	.	.	A	.	.	.	T	.	.	.	.	T	.	.	.	.	C	.
C. ignita haplotype H47	A	.	A	.	.	A	T	.	A	.	C	.	T	.	.	G	.	.
C. impressa haplotype H48	.	.	A	A	G	.	T	.	A	.	C	.	T	.	.	.	.	.
C. impressa haplotype H49	.	.	A	.	G	.	T	.	A	.	C	.	T	.	.	.	.	.

Figure 196.

Kimura-2-parameter neighbour joining tree of all Chrysis borealis sp. n. samples and haplotypes of C. ignita and C. impressa (sensu Soon et al. 2014). The C. terminata haplotype H163 (sensu Soon et al. 2014) is included as an outgroup.

Chrysis terminata Dahlbom, 1854

Figs 153, 154

Chrysis terminata Dahlbom, 1854: 261.

Chrysis ignita form A sensu Linsenmaier 1959: 156.

Diagnosis

Length 6–10 mm. The species resembles closely C. ignita by its colour, structure and habitus. However, the frontal carina has medially four tooth-like tubercles in both sexes (Figs 153, 154), the mesosoma is more homogeneously blue without extensive green areas on the pronotum and the mesoscutellum, and the body is slightly more elongate with more parallel sides.

Distribution

Latvia, Lithuania, Norway, Sweden. Relatively common (especially in south-eastern Sweden, Öland and Gotland). – West Palearctic: from western Europe to central Asia (Linsenmaier 1997).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood, but also sparsely vegetated sandy areas. Flight period: late April to early August. The flight begins earlier than in other species of the C. ignita group. Host: Ancistrocerus nigricornis (Curtis) (Vespidae) (van Lith 1954, Linsenmaier 1959, our own obs.).

Chrysis smaragdula group

Chrysis sexdentata Christ, 1791Fig. 81

Chrysis sexdentata Christ, 1791: 404.

Diagnosis

Length 7–11 mm. The species is easily recognised due to its unique combination of a red metasoma and six apical teeth. The head and the mesosoma are greenish, dark blue or nearly black with coppery reflections, whereas the metasoma is dorsally purple-red or coppery red. The tergites are coarsely punctured, and the posterior margin of T3 has six sharp teeth (Fig. 81). The shape of the body is robust and compact.

Distribution

Latvia. Very rare. Only one male specimen is known from central Latvia (Ropaži, 1.VI.1961, leg. V. Tumšs) (Tumšs and Maršakovs 1970). – Trans-Palearctic: from Europe and northern Africa to western and central Asia (Linsenmaier 1959, 1999).

Biology

Habitat: gardens with dead wood, old brick walls, old fences and/or stones (Trautmann 1927, 1930). Adults occasionally visit flowers of Apiaceae and Euphorbiaceae (Linsenmaier 1997, Rosa 2004). Flight period: June to July. Host: Euodynerus dantici (Rossi) (Martynova and Fateryga 2015) and possibly also Ancistrocerus parietum (Linnaeus) (Vespidae) (Mocsáry 1912, Berland and Bernard 1938). Host records implicating solitary bees (e.g. Osmia brevicornis (Fabricius), O. caerulescens (Linnaeus), Hoplitis adunca (Panzer) and Megachile sicula (Rossi)) (Trautmann 1927) are dubious, as pointed out by Kunz (1994).

Chrysis equestris Dahlbom, 1854

Figs 80, 89, 112, 134, 151

? Chrysis sexdentata Dahlbom, 1831: 30, not Christ, 1791.

Chrysis zetterstedti of authors, not Dahlbom, 1845.

Chrysis equestris Dahlbom, 1854: 307.

Chrysis fasciata of authors, not Olivier, 1790.

Diagnosis

Length 7–10 mm. Both sexes have a mostly dark blue or black, partially violet, body with green reflections on the frons, margins of the pronotum, mesoscutum, mesoscutellum and mesopleuron. The tergites have contrasting golden red or golden green bands posteriorly (except on the apical rim), which are especially wide laterally on T1 and T2. The colour and form of the bands is quite variable, usually they are wider and more reddish in the female than in the male. The species closely resembles C. zetterstedti, but is characterised by the following differences: the black spots of S2 are narrower, usually not extending to the lateral margins of the sternite (Fig. 112), T5 of the female (on ovipositor) is broader and has a longitudinal medial groove (Fig. 89), the head is broader, especially in female (shortest distance between the compound eyes is slightly longer than the diameter of an eye) (Fig. 151), the gonostyle is shorter, the cuspis is apically curved (not straight) (Fig. 134), and the propodeal tooth is slightly convex or straight ventrally (not lobate).

Distribution

Estonia, Finland, Lithuania, Norway, Sweden. Rare. – Trans-Palearctic: from western Europe to Russian Far East (Sakhalin).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood. Adults are usually found on sun-exposed dead tree trunks and stumps, most often of Populus, but also of Salix, Betula and Alnus, rarely Picea and Pinus. They also fly near log piles, telephone poles and walls of old wooden buildings (Frey 1915, Linsenmaier 1997). Flight period: mid-May to early August. Host: Discoelius dufourii Lepeletier and D. zonalis (Panzer) (Vespidae) (Pärn et al. 2014, our own obs.).

Chrysis zetterstedti Dahlbom, 1845

Figs 90, 113, 135, 152

? Chrysis sexdentata Dahlbom, 1831: 30, not Christ, 1791.

Chrysis Zetterstedti Dahlbom, 1845: 11.

Chrysis fasciata of authors, not Olivier, 1790.

Diagnosis

Length 6–9 mm. The species resembles C. equestris, but differs from it by the following characters: the black spots of S2 are broader, extending to the lateral and anterior margins of the sternite (Fig. 113), T5 of the female is narrower and does not have a longitudinal medial groove (Fig. 90), the head is narrower (the shortest distance between the compound eyes is shorter or as long as the diameter of an eye) (Fig. 152), the gonostyle is more elongated, as long as the cuspis, the cuspis is apically straight (not curved) (Fig. 135), and the propodeal tooth is weakly lobate ventrally (not convex or straight).

Distribution

Estonia, Latvia, Lithuania, Sweden. Rare. – Trans-Palearctic: from North Europe to Siberia. Records from the East Palearctic Region refer to C. fasciata daphne Smith, 1874 (Rosa et al. 2014).

Biology

Habitat: forest margins and clearings with sun-exposed dead tree trunks and stumps (e.g. Quercus). Flight period: probably similar to that of C. equestris, most specimens have been collected in July. Host: unknown, possibly Euodynerus notatus (Jurine) (N. Johansson pers. obs.).

Remarks

Several authors have considered C. zetterstedti to be either a synonym (e.g. Trautmann 1927, Kimsey and Bohart 1991) or a subspecies (Linsenmaier 1959, 1997, Rosa and Soon 2012) of C. fasciata. However, molecular and morphological studies have shown that C. zetterstedti most likely represents a valid species (Paukkunen et al. 2014). The occurrence of C. zetterstedti in central and southern Europe is still uncertain.

Genus Trichrysis Lichtenstein, 1876

Figs 197, 198–199

Trichrysis Lichtenstein, 1876: 27.

Note

In Europe, this genus is characterised by the tridentate posterior margin of T3 (Figs 197, 198), the medially located small black spots of S2 (Fig. 199), and the simple metanotum (Kimsey and Bohart 1991). The European species are relatively small (body length ca 4–8 mm) and completely blue or green in colour. Small males are often blackish, at least dorsally. The hosts are cavity-nesting crabronid and pompilid wasps, and possibly also solitary vespid wasps and megachilid bees (Trautmann 1927, Pärn et al. 2014). The genus includes 27 species in the Palearctic, Afrotropical and Oriental Regions (Kimsey and Bohart 1991, Strumia 2009). Three species have been found in Europe (Rosa and Soon 2012), and one, T. cyanea, from the Nordic and Baltic countries (Paukkunen et al. 2014).

Figure 197.

Trichrysis cyanea ♀. Scale 1 mm.

Figures 198–199.

Trichrysis cyanea ♀: 198 T3, dorsal view 199 S2, ventral view. Scale 1 mm.

Trichrysis cyanea (Linnaeus, 1758)

Figs 197, 198–199

Sphex cyanea Linnaeus, 1758: 572.

Trichrysis cyanea: Trautmann and Trautmann 1919: 34.

Diagnosis

Length 4–8 mm. The species is characterised by its completely green, blue or violet body (Fig. 197) and the tridentate posterior margin of T3 (Fig. 198). The male in particular is often dorsally or rarely almost completely black. The lateral teeth of T3 are often more like angles, and sometimes all teeth can be small or rounded and inconspicuous. The black spots of S2 are small and located close together (Fig. 199).

Distribution

Denmark, Estonia, Finland, Latvia, Lithuania, Norway, Sweden. Very common. – Trans-Palearctic: from Europe and northern Africa to central Asia, Siberia, Korea, China and Japan (Linsenmaier 1999, Kurzenko and Lelej 2007, Rosa et al. 2014).

Biology

Habitat: forest margins, clearings and gardens with sun-exposed dead wood (e.g. dead tree trunks, log and branch piles, walls of wooden buildings or poles). Adults occasionally visit flowers of Apiaceae and Rosaceae (Kusdas 1956, Rosa 2004, our own obs.). Flight period: late May to early September. Host: primarily species of Trypoxylon Latreille (Crabronidae), but also Auplopus carbonarius (Scopoli) and species of Dipogon Fox (Pompilidae), and possibly other cavity-nesting crabronid wasps (Dufour and Perris 1840, Morgan 1984, Brechtel 1986, Asís et al. 1994, Gathmann and Tscharntke 1999, Tormos et al. 1996, Pärn et al. 2014, our own obs.). Also many cavity-nesting solitary bee species have been reported as hosts, but these records are rather unreliable due to the very different biology of bees compared to crabronid and pompilid hosts.

Genus Chrysura Dahlbom, 1845

Figs 200, 201–208

Chrysura Dahlbom, 1845: 6.

Holochrysis Rye, 1878: 134.

Note

Diagnostic characters of this genus include the nearly flat and densely punctate frons, the lack of a transverse frontal carina (Figs 201, 202), and the long malar space. Usually the mandible is also toothed subapically (Fig. 202), the proximal flagellomeres of the male are swollen ventrally (Fig. 204) and the pronotum is shorter than the mesoscutellum. The radial cell of the forewing is closed and the posterior margin of T3 is rounded without apical teeth (Figs 206–208). Chrysura is the second largest genus in the tribe Chrysidini. It includes 117 valid species, of which 106 are distributed in the Palearctic Region (Kimsey and Bohart 1991, Rosa and Lotfalizadeh 2013). The hosts consist of solitary bees of the family Megachilidae. The European fauna includes 50 species and several subspecies (Rosa and Soon 2012). Five species have been recorded in the Nordic and Baltic countries (Paukkunen et al. 2014). Species-groups below follow Kimsey and Bohart (1991).

Figure 200.

Chrysura radians ♀. Scale 1 mm.

Figures 201–208.

Head, frontal view: 201 C. austriaca ♀ 202 C. hirsuta ♀. Metanotal tooth, dorsal view: 203 C. hirsuta ♀. Antenna: 204 C. radians ♂. Mesoscutum, mesoscutellum, metanotum and propodeum, lateral view (arrow indicating metascutellum): 205 C. trimaculata ♀. Metasoma, dorsal view: 206 C. trimaculata ♀, 207 C. radians ♀ 208 C. hirsuta ♀. Scale 1 mm.

Key to Chrysura species of the Nordic and Baltic countries

1	Mesosoma dorsally bright red	C. dichroa (Dahlbom)
–	Mesosoma entirely blue-green with golden (or rarely coppery) reflections, without red colour	2
2	Head broad ventrally, with nearly parallel malar spaces, width between mandible bases about 1.5 times as long as compound eye (Fig. 201). Mandibles without or with very small subapical tooth. Male with simple antennal segments. Metascutellum evenly rounded, not elevated medially	C. austriaca (Fabricius)
–	Head narrow ventrally, with convergent malar spaces, width between the mandible bases equivalent or slightly longer than length of compound eye (Figs 202). Mandibles with large subapical tooth (Fig. 202). Male with ventrally swollen antennal segments (Fig. 204). Metascutellum medially elevated or relatively rounded	3
3	T3 of female long and ovoid, posterior pit row very weakly developed (Fig. 206). Metascutellum sharply elevated medially (Fig. 205). Punctation of tergites very dense and homogeneous (Fig. 206). Setae on posterior margin of T3 brown	C. trimaculata (Förster)
–	T3 of female short and wide, with distinct pit row posteriorly (Figs 207, 208). Metascutellum relatively rounded, not sharply elevated medially. Punctation of tergites homogeneous or heterogeneous (Figs 207, 208). Setae on posterior margin T3 brown or white	4
4	Punctation of tergites heterogeneous, consisting of large punctures and interspersed small punctures (Fig. 207). Setae on T3 white. Metanotal tooth indistinct. Third antennal segment about 3.5 times as long as broad	C. radians (Harris)
–	Punctation of tergites homogeneous, consisting of only small punctures (Fig. 208). Setae on T3 brown. Metanotal tooth distinct (Fig. 203). Third antennal segment about 3 times as long as broad	C. hirsuta (Gerstaecker)

Chrysura austriaca group

Chrysura austriaca (Fabricius, 1804)

Fig. 201

Chrysis austriaca Fabricius, 1804: 173.

Chrysura austriaca: Dahlbom 1845: 6.

Diagnosis

Length 8–12 mm. The species differs from other similarly coloured species of Chrysura by its ventrally broader head (Fig. 201) and simple mandible, which lacks or has only a very small subapical tooth (Fig. 201). The antennal segments of the male are not ventrally swollen. The head and mesosoma are blue, often with golden green reflections laterally on the mesoscutum, and the metasoma is golden red.

Distribution

Lithuania. Very rare. The species is known from three localities in Lithuania (Puvočiai, Trakai, Vilnius) (Orlovskytė et al. 2010). – Trans-Palearctic: from Europe and northern Africa to Siberia and Japan (Linsenmaier 1997, Kurzenko and Lelej 2007).

Biology

Habitat: forest margins and gardens. Often found from wooden poles, walls, fence posts, loess, clay or old brick walls or rocks (Trautmann 1927, Kunz 1994, Linsenmaier 1997). Adults occasionally visit flowers of Apiaceae and Euphorbiaceae(Linsenmaier 1997, Rosa 2004). Flight period: June to early August. In Lithuania, specimens have been collected in July and early August. Host: Hoplitis adunca (Panzer) (Megachilidae) (e.g. Frey-Gessner 1887, Trautmann and Trautmann 1919, Linsenmaier 1959), possibly also H. anthocopoides (Schenck) and Osmia parietina Curtis (Schenck 1856, Mocsáry 1879, Banaszak 1980).

Chrysura dichroa group

Chrysura dichroa (Dahlbom, 1854)

Chrysura nitidula Dahlbom, 1845: 7, nomen oblitum, not Chrysis nitidula Fabricius, 1775.

Chrysis Gyllenhali Dahlbom, 1854: 143.

Chrysis dichroa Dahlbom, 1854: 146.

Chrysura dichroa: Kimsey and Bohart 1991: 488.

Diagnosis

Length 5–9 mm. The species is easy to differentiate from other North European Chrysura species by its bright red pronotum, mesoscutum and mesoscutellum. The head, mesopleuron, metanotum, propodeum and legs (excluding tarsi) are green or blue, and the metasoma is golden red, as in other species of the genus. The tergites are very densely and finely punctured.

Distribution

Sweden. Very rare. Only one female specimen is known from Västergötland, southern Sweden, collected probably in the 1830s (leg. L. Gyllenhal). – West Palearctic: southern and central Europe, south-eastern Asia (Linsenmaier 1997).

Biology

Habitat: rock mounds, scree formations, rocky outcrops, rock walls and dry meadows, usually in areas with calcareous bedrock (Kunz 1994, Linsenmaier 1997). Adults occasionally visit flowers of Apiaceae, Asteraceae and Rosaceae (Linsenmaier 1997, Rosa 2004, 2006). Flight period: late May to mid-August. Host: species of Osmia Panzer (Megachilidae) which build nests in empty snail shells, primarily O. rufohirta (du Buysson 1891, Ferton 1905, Malyshev 1968, Bonelli 1974), but also O. aurulenta (Panzer), O. versicolor Latreille, O. andrenoides Spinola, O. spinulosa (Kirby), O. ferruginea Latreille and O. caerulescens (Linnaeus) (Dalla Torre 1892, Ferton 1905, Grandi 1959, Heinrich 1964).

Chrysura radians group

Genus Parnopes Latreille, 1797

Fig. 209

Parnopes Latreille, 1797: 126.

Note

The genus can be distinguished from other genera of Parnopini by the reduced palpi, the large metascutellar projection and the larger body size. Members of the genus are parasites of ground-nesting solitary wasps of the tribe Bembicini (Crabronidae: Bembicinae). A total of 16 species are recognised, most of which occur in the Palearctic and Nearctic Regions (with 4 and 7 species, respectively) (Kimsey and Bohart 1991). A few species are known from India and Africa. Only one species, P. grandior, is found in Europe (Rosa and Soon 2012). The genus has been divided into species-groups by Kimsey and Bohart (1991).

Parnopes grandior (Pallas, 1771)

Fig. 209

Chrysis grandior Pallas, 1771: 474.

Parnopes grandior: Mocsáry 1882: 74.

Diagnosis

Length 8–12 mm. The species is easy to differentiate from other North European cuckoo wasps by its unique structure and colouration. The head, mesosoma and most of T1 are green or green-blue, with golden or coppery reflections, especially in the female. The metasoma behind T1, tegulae and tibiae are usually non-metallic red (Fig. 209). Sometimes also T2, T3 (and T4) have a metallic sheen laterally or even dorsally. The female has three and the male four external tergites, and the posterior margin of the last tergite is irregularly dentate (Fig. 209). The mouthparts are longer than the rest of the head, and the tegulae are large covering the wing bases (Fig. 209).

Distribution

Lithuania. Very rare. The species has been recorded from five localities in southern Lithuania (Orlovskytė et al. 2010). – West Palearctic: from Europe and northern Africa to Yemen and southwestern Asia (Linsenmaier 1997, 1999).

Biology

Habitat: xerothermic sparsely vegetated sandy areas. Adults often visit flowers of several different families (Bischoff 1923, Trautmann 1927, Molitor 1935, Schneid 1954, Linsenmaier 1997, Rosa 2004). Flight period: July to August. Host: Bembix rostrata (Linnaeus) (Crabronidae) (Gauss 1967), in southern Europe also other species of Bembix Fabricius (Linsenmaier 1968, Kunz 1994). The larva does not consume the host larva until it is fully grown (Malyshev 1968).

Figure 209.

Parnopes grandior ♂. Scale 1 mm.

Table 3.

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Distribution of chrysidid species in the Nordic and Baltic countries. DK = Denmark, EE = Estonia, FI = Finland, LV = Latvia, LT = Lithuania, NO = Norway, SE = Sweden. 1 = recorded before 2000, 2 = recorded from 2000 onwards.

Species	DK	EE	FI	LV	LT	NO	SE
Cleptes nitidulus (Fabricius, 1793)	1, 2	2	1, 2	1			1, 2
Cleptes semicyaneus Tournier, 1879	1					2	2
Cleptes semiauratus (Linnaeus, 1761)	1, 2	1, 2	1, 2	1	1	1, 2	1, 2
Omalus biaccinctus (du Buysson, 1892)	1	1, 2	1, 2	1		2	1, 2
Omalus aeneus (Fabricius, 1787)	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Omalus puncticollis (Mocsáry, 1887)						2	2
Pseudomalus pusillus (Fabricius, 1804)	1			1, 2	1, 2
Pseudomalus auratus (Linnaeus, 1758)	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Pseudomalus triangulifer (Abeille de Perrin, 1877)	1	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Pseudomalus violaceus (Scopoli, 1763)	1, 2	1, 2	1, 2	1, 2	1	1, 2	1, 2
Philoctetes truncatus (Dahlbom, 1831)	1	1, 2		2			1
Elampus constrictus (Förster, 1853)	1	1	1, 2			1, 2	1, 2
Elampus foveatus (Mocsáry, 1914)		2	1, 2			2	1, 2
Elampus panzeri (Fabricius, 1804)	1, 2	1, 2	1, 2	1	1	1, 2	1, 2
Holopyga fervida (Fabricius, 1781)	1
Holopyga metallica (Dahlbom, 1854)			1, 2
Holopyga generosa (Förster, 1853)	1	2	1, 2	1	1, 2		1, 2
Holopyga inflammata (Förster, 1853)			1		1
Hedychrum gerstaeckeri Chevrier, 1869	1	1, 2	1, 2	1, 2	1, 2
Hedychrum rutilans Dahlbom, 1854	1, 2	1, 2	1, 2	1, 2	1, 2
Hedychrum nobile (Scopoli, 1763)	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Hedychrum niemelai Linsenmaier, 1959	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Hedychrum chalybaeum Dahlbom, 1854				1	1
Hedychridium zelleri (Dahlbom, 1845)			1, 2
Hedychridium ardens (Coquebert, 1801)	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Hedychridium coriaceum (Dahlbom, 1854)	1	2	1, 2	1, 2	1, 2		1, 2
Hedychridium cupreum (Dahlbom, 1845)	1, 2	1, 2	1, 2	1, 2	1	1, 2	1, 2
Hedychridium purpurascens (Dahlbom, 1854)		2
Hedychridium caputaureum Trautmann & Trautmann, 1919		2	1, 2	2	2		1, 2
Hedychridium roseum (Rossi, 1790)	1	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Pseudospinolia neglecta (Shuckard, 1837)	1, 2	1, 2	1, 2	1, 2	1, 2	1	1, 2
Spinolia unicolor (Dahlbom, 1831)	1			1			1, 2
Chrysis gracillima Förster, 1853		2
Chrysis bicolor Lepeletier, 1806	1, 2	1, 2	1, 2	1, 2	1		1, 2
Chrysis westerlundi Trautmann, 1927			1, 2
Chrysis illigeri Wesmael, 1839	1, 2	1, 2	1, 2	1	1, 2	1, 2	1, 2
Chrysis succincta Linnaeus, 1767	1			1, 2	1
Chrysis leachii Shuckard, 1837	1
Chrysis scutellaris Fabricius, 1794	1				1		1, 2
Chrysis splendidula Rossi, 1790				1, 2
Chrysis rutilans Olivier, 1791		1, 2	1, 2	1	1		1, 2
Chrysis pulcherrima Lepeletier, 1806	1
Chrysis viridula Linnaeus, 1761	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Chrysis graelsii Guérin-Méneville, 1842		1, 2	1, 2	1, 2	1, 2
Chrysis indigotea Dufour & Perris, 1840							1
Chrysis fulgida Linnaeus, 1761	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Chrysis iris Christ, 1791	1	1, 2	1, 2	1, 2	1, 2		1, 2
Chrysis ruddii Schuckard, 1837	1, 2	1, 2	1, 2	1, 2	1	1, 2	1, 2
Chrysis corusca Valkeila, 1971		1, 2	2	1	1, 2	2	1, 2
Chrysis clarinicollis Linsenmaier, 1951		2
Chrysis vanlithi Linsenmaier, 1959	1, 2					2	1, 2
Chrysis subcoriacea Linsenmaier, 1959	1	1, 2	1, 2	1		1, 2	1, 2
Chrysis angustula Schenck, 1856	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Chrysis longula Abeille de Perrin, 1879	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Chrysis brevitarsis Thomson, 1870		1, 2	1, 2		1		1, 2
Chrysis pseudobrevitarsis Linsenmaier, 1951	1	1, 2	1, 2	1	1, 2	2	1, 2
Chrysis mediata Linsenmaier, 1951	2	1, 2		1	1, 2		1, 2
Chrysis solida Haupt, 1957	1	1, 2	1, 2	1	1, 2	1, 2	1, 2
Chrysis leptomandibularis Niehuis, 2000		1, 2	1	1	1, 2	2
Chrysis schencki Linsenmaier, 1968	1, 2	1, 2	1, 2	1	1, 2	1, 2	1, 2
Chrysis ignita (Linnaeus, 1758)	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Chrysis impressa Schenck, 1856	1	1, 2	1, 2	1	1, 2	1, 2	1, 2
Chrysis borealis Paukkunen, Ødegaard & Soon, sp. n.	1	1, 2	1, 2			1, 2	1, 2
Chrysis terminata Dahlbom, 1854				1	1, 2	1, 2	1, 2
Chrysis sexdentata Christ, 1791				1
Chrysis equestris Dahlbom, 1854		1, 2	1, 2		1, 2	1, 2	1, 2
Chrysis zetterstedti Dahlbom, 1845		1, 2		1	1, 2		1, 2
Trichrysis cyanea (Linnaeus, 1758)	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2	1, 2
Chrysura austriaca (Fabricius, 1804)					1, 2
Chrysura dichroa (Dahlbom, 1854)							1
Chrysura hirsuta (Gerstaecker, 1869)		1, 2	1, 2			1, 2	1, 2
Chrysura radians (Harris, 1776)	1	1, 2		1	1, 2	1, 2	1, 2
Chrysura trimaculata (Förster, 1853)							1, 2
Parnopes grandior (Pallas, 1771)					1, 2
Number of species	47	51	48	49	48	38	53

Acknowledgements

We thank Johan Abenius (Sweden), Niklas Johansson (Sweden), Kent Runge Poulsen (Denmark) and Hans Th. Schmidt (Denmark) for providing important information and help for this study. Pekka Malinen (Finland) and Arnstein Staverløkk (Norway) gave permission to use their photographs. We also thank Mattias Forshage (Sweden) and Lauri Kaila (Finland) and the reviewers for their helpful comments on the manuscript and Stephen Venn (Finland) and Robert Davis (Estonia) for checking the English language. This research was supported by the Finnish Ministry of the Environment through the research programme of deficiently known and threatened forest species (PUTTE II), the institutional grant IUT20-33 from the Estonian Research Council, the European Union through the European Regional Development Fund (Center of Excellence FIBIR) and the project ACUNOR as a part of the Taxonomy Initiative of the Norwegian Biodiversity Information Centre. DNA barcode data used in this publication was generated in collaboration with the Finnish Barcode of Life project (FinBOL) funded by the Kone Foundation and the University of Oulu, and the Norwegian Barcode of Life Network (NorBOL) funded by the Research Council of Norway and the Norwegian Biodiversity Information Centre. Societas Entomologica Helsingforsiensis also gave support for DNA barcoding, and the Entomological Society of Lund (ESIL) gave funds for acquiring the imaging system.

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