Biodiversity Data Journal :
Taxonomy & Inventories
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Corresponding author: Jeremy Joshua Hübner (huebner@snsb.de), Vasilisa Chemyreva (diapriidas.vas@gmail.com)
Academic editor: Ralph Peters
Received: 20 Oct 2023 | Accepted: 20 Dec 2023 | Published: 08 Jan 2024
© 2024 Jeremy Hübner, Vasilisa Chemyreva
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:
Hübner JJ, Chemyreva V (2024) Review of German Spilomicrus Westwood (Hymenoptera, Diapriidae, Spilomicrini). Biodiversity Data Journal 12: e114515. https://doi.org/10.3897/BDJ.12.e114515
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This study provides an integrative taxonomy-based review for the genus Spilomicrus Westwood in Germany using DNA barcoding and classic morphology.
Spilomicrus simplex Tomsik, 1947 is placed in synonymy with S. antennatus Jurine, 1807; Spilomicrus thomsoni Kieffer, 1911 is removed from synonymy with S. hemipterus Marshall, 1868. A lectotype is designated for Spilomicrus nigripes Thomson, 1858. Newly recorded for Germany are the following species: S. thomsoni Kieffer, 1911, S. crassiclavis Marshall, 1868, S. lusitanicus Kieffer, 1910 and S. diversus Chemyreva, 2021. Three species, Spilomicrus brevimalaris sp. nov., S. flavecorpus sp. nov. and S. politus sp. nov. are described as new to science. The 23 DNA-barcodes with species identification present a substantial addition over the previous German checklist. This study aims to update the number of nationwide known Spilomicrus species from fifteen to twenty. Furthermore, a new key to identify all European Spilomicrus species is provided.
checklist, DNA-barcoding, integrative taxonomy, key, new records, new species, new synonymy, parasitoid wasps
This study provides a review of the diaprid genus Spilomicrus (Diapriinae, Spilomicrini) in Germany. Diapriidae are parasitoid wasps that are referred to as a “dark taxon” because they are hyper-diverse and it is assumed that a large proportion of the species diversity remains hidden (
The most recent diversity evaluation that was conducted for Germany was done so over twenty years ago by
Overall, the German diaprid fauna is expected to resemble the European species communities which have been recently examined in detail by
In order to tackle this megadiverse “dark taxon”, we take advantage of DNA barcoding (
Most of the material was collected in Germany in various collecting events, mainly in Bavaria in the framework of GBOL III: Dark Taxa project. Part of the investigated specimens were taken from the Hilpert collection. All specimens are stored at the Bavarian State Collection of Zoology in Munich. In addition, type material from various museums was examined. For species identification, we applied an integrative taxonomy approach, using all resources possible: barcoded and non-barcoded material, as well as genetic and morphological identification methods. Based on the CO1 barcodes which we obtained from the Canadian Centre for DNA Barcoding (CCDB, https://ccdb.ca/resources/), a Maximum-Likelihood tree was calculated using IQ TREE (online tool,
The morphological terminology and abbreviations follow Hymenoptera Anatomy ontology (
The following abbreviations for locations in Germany are used: BW = Baden-Wuerttemberg, BY = Bavaria, HE = Hesse, NRW = North Rhine-Westphalia. Museum acronyms: HNHM – Hungarian Natural History Museum, Budapest, Hungary; MNHN – National Museum of Natural History, Paris, France; MZLU – Lund Museum of Zoology, Sweden; NHRS – Swedish Museum of Natural History, Stockholm, Sweden; MMBC – Moravian Museum, Brno, Czech Republic; ZISP – Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia.
Type species Spilomicrus stigmaticalis Westwood, 1832, by original monotypy.
A detailed diagnosis of the genus was given by
Medium-sized (1.5–4.5 mm long) melanic wasps. Head subglobose, with mouthparts in lateral view hypognathous; antenna 13-segmented, in females with clava more- or less abrupt, in males antenna thread-like, A4 modified in almost all species. Mesosoma moderately to distinctly wider than high; scutellum with 2 anterior pits and, in most species, with 2 lateral pits and row of smaller posterior pits along posterior margin; forewing with costal vein tubular to nebulous, submarginal vein tubular, marginal vein relatively short, postmarginal and stigmal veins rudimentary or absent; basal vein rarely tubular, in most species nebulous or absent; other veins, at most, nebulous or absent; legs slender to stout, with or without trochanters. Petiole cylindrical in most species; anterior margin of T2 straight, without median cleft or emargination (rarely with 2 lateral folds filled with pilosity); base of S2 arcuate, with moderate to strong cushion of pilosity.
The following part lists all the Spilomicrus species found within the framework of the GBOL III project. In comparison to the whole European Spilomicrus fauna, three species could not be recorded for Germany and are, therefore, not documented here: S. sanbornei Masner, 1991, S. cursor Kieffer, 1911 and S. latus Chemyreva, 2021. In addition to the morphology, we provide the barcoding information in the form of the BINs and, if necessary, genetic distances for closely-associated taxa. Illustrations are given for the newly-described taxa and the closest sister taxa for a better understanding of the morphological characters and differences. All other species have already been well described and illustrated in
Spilomicrus abnormis Marshall, 1868 : 202.
Spilomicrus minimus Kieffer, 1911. Synonymised by
Illustrated in
Czech Republic, Germany*, Hungary, Ireland, Korea, Moldova, Netherlands, Poland, Russia.
Spilomicrus annulicornis Kieffer, 1911 : 788.
Illustrated in
Austria, Czech Republic, Finland, France, Germany, Netherlands, Russia (European part), United Kingdom.
Psilus antennatus Jurine, 1807 : 319.
Basalys californica Ashmead, 1893. Synonymised by
Eriopria nigra Kieffer, 1910. Synonymised by
Eriopria rufithorax Kieffer, 1910. Synonymised by
Scutellipria quinquepunctata Szabo, 1961. Synonymised by
Spilomicrus simplex Tomsik, 1947 : 33, 34, 40. Syn. nov. Fig.
Austria, Bulgaria, Czech Republic, Germany, Hungary, Romania, Slovakia, Switzerland, United Kingdom, United States.
What was already suspected by some researchers could be established using DNA barcoding of specimens of each sex (only one female was available, but numerous males). Our obtained sequences were too short to be included in the attached tree (Suppl. material
Spilomicrus bipunctatus Kieffer, 1911 : 284, 289.
Illustrated in
Azerbaijan, Czech Republic, Estonia, France, Germany, Hungary, Italy, Moldova, Netherlands, Poland, Russia (European part), Slovakia, Ukraine, United Kingdom.
Male. Body length 1.4 mm; forewings reaching far beyond apex of metasoma; antenna 0.9 times as long as body.
Head: black; in dorsal view 1.35 times as wide as long, as wide as mesosoma. Temples behind eyes gradually receding posteriorly. Tentorial pit tiny. Malar sulcus absent. Clypeus weakly convex, oval, 1.7 times as wide as high. Mandible reddish-brown, elongate, with upper tooth slightly shorter than lower tooth. Palpi yellow. Eye oval, with scattered long setae; 0.6 times as high as head and 3.8 times as high as malar space. Frons above base of toruli smooth. Postgenal cushion scanty (Fig.
Male holotype Spilomicrus brevimalaris sp. nov. (ZSM-HYM-33100-G04; BOLD:AEC2138). A lateral; B dorsal; C antenna.
Female. Body length 1.6–1.7 mm. Wings 0.9–1.0 times as long as the body. Pleurostomal distance 0.8 times as long as shortest distance between eyes (Fig.
Female paratype Spilomicrus brevimalaris sp. nov. (ZSM-HYM-33108-G09; BOLD:AEC2138). A lateral; B dorsal; C face frontal; D antenna.
Male. Body length 1.3–2.1 mm. Face without malar sulcus, pleurostomal distance slightly wider than shortest distance between eyes (Fig.
The name of this species is a composite Latin masculine adjective derived from “brevis” and “malar” and refers to the short malar distance typical for the males of the new species.
Germany, Russia (European part). Further BIN records are online available for Italy and Norway. Probably further distributed around western Europe.
The male specimen was used in this case as a holotype, since there is no possibility to use females for the S. lusitanicus-species group (both species, S. brevimalaris sp. nov. and S. flavecorpus sp. nov., are very close to S. lusitanicus (Kieffer)). There are two reasons for that: 1) the female for the S. lusitanicus is unknown; 2) The most reliable feature to determine this species is the length of the malar space, but this feature does not work for the female determination.
The Russian material that was recorded by
Spilomicrus compressus Thomson, 1859 : 369.
Spilomicrus carinatus Kieffer, 1911. Synonymised by
Spilomicrus crassipes Kieffer, 1911. Synonymised by
Illustrated in
Belarus, Czech Republic, Estonia, France, Germany, Hungary, Poland, Russia (European part), Sweden, Ukraine, United Kingdom.
Spilomicrus crassiclavis Kieffer, 1911 : 788, 797.
Spilomicrus pelion Nixon, 1980. Synonymised by
Illustrated in
Czech Republic, Denmark, Finland, Germany*, Japan, Norway, Sweden, United Kingdom.
Spilomicrus diversus Chemyreva, 2021 : 19.
Female. Face with malar sulcus visible in the form of shallow furrow. Malar distance 0.47 times as long as largest diameter of eye. Front behind scapus with two small holes (as in the male, Fig.
Male Spilomicrus diversus (ZSM-HYM-42367-C03; BOLD:ADF4749). A lateral; B dorsal; C head dorsofrontal, small oval holes marked with red arrow; D antenna.
Female Spilomicrus diversus (ZSM-HYM-42318-D01; BOLD:ADF4749). A lateral; B dorsal; C face; D antenna dorsal.
Male. Antennae filiform (Fig.
Abkhazia, Czech Republic, Georgia, Germany*, Poland, Russia (European part: Samara Prov., Republic Adygea; Siberia: Krasnoyarskiy Terr.).
Based on new data on intraspecific variability of the Spilomicrus diversus and re-investigation of the type series, we conclude that some specimens should be excluded from the type series. The front sculpture of the specimens from the Far East of Russia (Primorskiy Terr. and Sakhalin Area) is significantly different from both S. diversus and S. politus sp. nov. and these specimens (paratypes) must be excluded from the type series.
Male. Body length 1.9 mm; forewings reaching far beyond apex of metasoma; antenna 0.8 times as long as body.
Head: brown; in dorsal view 1.05 times as wide as long, as wide as mesosoma. Temples behind eyes gradually receding posteriorly. Tentorial pit tiny. Malar sulcus absent. Clypeus weakly convex, oval, 1.85 times as wide as high. Mandible brown, elongate, with upper tooth shorter than lower tooth. Palpi yellow. Eye oval, with few scattered long setae; 0.4 times as high as head and 1.7 times as high as malar space. Frons above base of toruli smooth. Postgenal cushion scanty (Fig.
Female. Body length 1.7–1.8 mm. Pleurostomal distance 0.74 times as long as shortest distance between eyes (Fig.
Male. Face without malar sulcus, pleurostomal distance 0.9 times as wide as shortest distance between eyes (Fig.
The name of this species is a composite Latin masculine adjective derived from the adverb “flave” (yellowly) and “corpus” and refers to the colouration of the body.
Germany. Further BIN records are online available for Canada. Probably further distributed around the Palaearctic and Nearctic.
The reason for the selection of the holotype is analogous to that of S. brevimalaris sp. nov. (check Notes).
Spilomicrus flavipes Thomson, 1858: 369.
Spilomicrus szelenyii Szabo, 1977. Synonymised by
Illustrated in
Czech Republic, France, Germany, Hungary, Ireland, Moldova, Mongolia, Poland, Russia, Sweden, United Kingdom.
Spilomicrus formosus Jansson, 1942 : 215.
Illustrated in
Belgium, Canada, Czech Republic, Denmark, Finland, Germany, Great Britain, Ireland, Japan, Norway, Russia, Slovakia, Sweden, United States.
Spilomicrus hemipterus Marshall, 1868 : 202.
Spilomicrus inaequalis Tomsik, 1941: 34, 38, 42. Synonymised by
Spilomicrus pedisequus Kieffer, 1916: 784, 787. Synonymised by
Malar sulcus partly developed, shallow; neck of prothorax bare anteriorly; pleurostomal distance distinctly shorter than distance between eyes in front view; temples behind eyes gradually receding posteriorly; male А4 with keel reaching 0.7 of the segment length, A4 0.65 times as long as A3, widened apically; antenna distinctly bicolorous with more abrupt clava; female A9 without or with weakly indicated MGS brush; A13 with distinct small pit ventrally; A13 in dorsal and lateral views narrower than A12; А9 distinctly narrower and shorter than А10; notauli present in the form of broad grooves posteriorly; sternaulus absent; wings reaching to one-fourth of metasoma length to distinctly beyond the apex of metasoma; female petiole elongate, about 1.2 times as long as wide.
Austria, Croatia, Czech Republic, France, Germany, Hungary, Moldova, Netherlands, Poland, Russia (European part), Switzerland, Ukraine, United Kingdom.
Spilomicrus integer Thomson, 1859 : 369.
Spilomicrus major Vollenhoven, 1879. Synonymised by
Illustrated in
Czech Republic, France, Germany, Hungary, Netherlands, Poland, Romania, Russia (European part), Slovakia, Sweden, Ukraine, United Kingdom.
Tritopria lusitanica Kieffer, 1910 : 749, male. Fig.
Male types. A holotype of Spilomicrus noctiger Szabó; B corresponding labels; D, F lectotype of Spilomicrus gracilicornis Kieffer (designated by
Spilomicrus gracilicornis Kieffer, 1911. Synonymised by
Spilomicrus noctiger Szabo, 1977. Synonymised by
Male. Body length 1.9–2.5 mm. Face without malar sulcus, pleurostomal distance slightly wider than shortest distance between eyes. Malar distance 0.45–0.55 times as long as largest diameter of eye. Front smooth. Аntennae dark brown, slender and long, with A5–A12 2.3–3.3 times as long as wide. A4 1.2–1.25 times as long as A3 and with keel and emargination reaching to 0.5–0.55 of the segment length (Fig.
Male Spilomicrus lusitanicus (ZSM-HYM-42423-H01; BOLD:AEK2205). A lateral; B dorsal; C antenna lateral; D antenna dorsal.
Algeria, Austria, Czech Republic, France, Germany*, Hungary, Italy*, Portugal, Russia (European part).
The most important features of this species, such as malar and pleurostomal distances cannot be examined in the lectotype of Tritopria lusitanica because the face of the type specimen is hidden in glue. However, secondary diagnostic characters (proportions of the remaining antennomeres, width of the head and proportions of the scutellum) lead us to believe that all type specimens belong to a single species and correspond with the examined material mentioned above under the name Spilomicrus lusitanicus. The females are unknown. The females described by
Spilomicrus modestus Tomsik, 1947 : 33, 39, 42.
Illustrated in
Austria, Czech Republic, Finland, Germany, Hungary, Moldova, Russia (European part and East Siberia), Ukraine.
Spilomicrus nigriclavis Marshall, 1868 : 228.
Spilomicrus punctatus Kozlov, 1978 : 591, nom. praeocc., non Spilomicrus punctatus (Cameron, 1889).
Spilomicrus kozlovi Notton, 2014. Synonymised by
Spilomicrus nigriclavis var. armatus Kieffer, 1911 : 781, nom. praeocc., non Spilomicrus armatus (Ashmead, 1893).
Spilomicrus nigriclavis var. subarmatus Kieffer, 1912. Synonymised by
Illustrated in
France, Germany, Netherlands, Russia (European part), Sweden, United Kingdom.
Female (holotype). Body length 1.8 mm; forewing extending far beyond apex of metasoma; antenna 0.68 times as long as body. Head: black, in dorsal 0.95 times as wide as metasoma. Tentorial pits absent. Clypeus weakly convex, 0.6 times as high as wide. Mandible dark brown, elongate, its upper tooth slightly shorter than lower tooth. Palpi yellow. Eye oval, with scattered long setae, 0.42 times as high as head and 1.9 times as high as malar space. Postgenal cushion dense. Antennae: A1 slightly curved, broadened apically, finely coriaceous; its apical rim simple. A2 not compressed. Apical half of A1 and A2–A8 dark brown, A9–A13 dark brown. Antenna A10–A13 with MGS brush, flattened on ventral side. A10–A12 as long as wide. A13 distinctly narrower than A12 and 1.1 times as long as A12. Antennomers length to width ratios in dorsal view as in Fig.
Male (BOLD: AER1505). Body length 1.6 mm. Similar to female, but differs by the following features: antenna filiform, A2–A13 brown, A1 dark brown (Fig.
Male Spilomicrus politus sp. nov. (ZSM-HYM-42318-B01; BOLD:AER1505). A lateral; B dorsal; C head dorsal without holes, bare area marked with red arrow; D antenna.
The species closely resembles S. diversus Chemyreva, 2021 from which it can be distinguished by the combination of the following features: A11 and A12 2.7 times as wide as A5 (A11–A12 about 2.3 times as wide as A5 in S. diversus); the malar sulcus is totally absent (visible in the form of shallow furrow in S. diversus); frons above base of toruli smooth (Fig.
The name of the new species is a Latin masculine adjective “politus” (smooth).
Estonia, Georgia (Republic of Abkhazia and Autonomous Republic of Adjara), Germany, Romania, Russia (European part).
The new species Spilomicrus politus sp. nov. was assigned two BINs, BOLD:ACZ2358 and BOLD:AER1505. It was not reliably possible to separate those two BINs into two morphologically sound species. The distance between those two BINs is 1.74%, whereas the distances to Spilomicrus diversus (BOLD:ADF4749) are 2.59% (BOLD:ACZ2358) and 3.12 % (BOLD:AER1505), the distance to S. modestus is 13.6%. The fact that both BINs of the S. politus sp. nov. differ in under 2% of the bases in their sequences leads to the suspicion that the specimens might just be one species.
Spilomicrus rufitarsis Kieffer, 1911 : 786.
Spilomicrus pseudocursor Szabo, 1974 : 497. Synonymised by
Illustrated in
Algeria, Austria, Czech Republic, France, Germany, Hungary, Ireland, Italy, Netherlands, United Kingdom.
Spilomicrus stigmaticalis Westwood, 1832 : 129, female.
Spilomicrus nigripes Thomson, 1859. Synonymised by
Spilomicrus basalyformis Marshall, 1868. Synonymised by
Spilomicrus armatus Ashmead, 1893. Synonymised by
Spilomicrus tripartitus Kieffer, 1911. Synonymised by
Spilomicrus pilicornis Szabo, 1977b. Synonymised by
Spilomicrus barbatus Szabo, 1983. Synonymised by
Spilomicrus mediofurcatus Szabo, 1983. Synonymised by
Algeria, Azerbaijan, Austria, Canada, Czech Republic, Finland, France, Georgia, Germany, Hungary, Ireland, Italy, Kazakhstan, Moldova, Netherlands, Poland, Russia (European part and Siberia), Slovakia, Sweden, Switzerland, Ukranie, United Kingdom, United States.
Spilomicrus stigmaticalis is a fairly common, widely distributed species. The species contains two BINs, BOLD:ADS1706 and BOLD:ACU1243. Still, all sequences are clustered as one single taxon using the BOLD cluster analysis and the ASAP algorithm. Not only is the genetic distance between those BINs small (1.9%), they also show medium to high intraspecific variation of up to 2.2% (mean distance 0.6%). In addition to that, we were not able to distinguish both genetic clades morphologically in both sexes, not even using the genitalia. It was only possible to find identifying morphological characters to distinguish between the females. Due to the genetic and morphological proximity of both clades, we will keep them together as one species. A lectotype is designated for Spilomicrus nigripes Thomson, 1858 (Fig.
Spilomicrus thomsoni Kieffer, 1911 : 787, 798.
Malar sulcus partly developed, shallow; neck of prothorax bare anteriorly; pleurostomal distance distinctly shorter than distance between eyes in front view; temples behind eyes gradually receding posteriorly; male А4 cylindrical, with keel reaching 0.55 of the segment length, A4 0.73–0.80 times as long as A3; antenna gradually darkened towards the top, with non-abrupt clava; female A9 with distinct MGS brush; A13 with small pit ventrally; A13 in dorsal and lateral views narrower than A12; А9 distinctly narrower and shorter than А10; notauli present in the form of broad grooves posteriorly; sternaulus absent; wings reaching to apex of metasoma to distinctly beyond it; female petiole elongate, about 1.2 times as long as wide. Lectotype illustrated in Fig.
Czech Republic (Tomsik 1947), Finland, Germany*, Moldova, Russia (European part), Sweden, Ukraine.
There are two BINs within Spilomicrus thomsoni, BOLD:ADF4747 and BOLD:ADX1651, which differ in only 0.1% from each other. Although the cluster methods of ASAP and BOLD separate the two clades and show very low intraspecific genetic variation, we could not tell them morphologically apart. Therefore, we will refer to them as being one species until further analyses might change that interpretation.
On the other hand, we can separate the Spilomicrus thomsoni taxon from S. hemipterus genetically and morphologically. This is why we removed S. thomsoni from synonymy with S. hemipterus.
Key to the European Spilomicrus species (modified and updated after Chemyreva (2021)) Females (female of S. lusitanicus unknown) |
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1 | Ventral margin of clypeus with pointed or rounded deflexed median projection; mandibles short, with upper tooth much shorter than lower tooth | 2 |
– | Ventral margin of clypeus with rounded reflexed median projection (fig. 17, 2 [arrow], 4 in |
4 |
2 | Antennae with clava 5- or 6-segmented; in front view, ventral margin of clypeus rounded, blunt; mesosoma depressed, no more than 0.8 times as high as wide, mesoscutum weakly convex; median propodeal keel low, hardly raised anteriorly | S. sanbornei |
– | Antennae with clava 7- or 8-segmented; in front view, ventral margin of clypeus triangular, pointed; mesosoma less depressed, at least 0.9 times as high as wide, mesoscutum strongly convex; median propodeal keel distinctly raised anteriorly to form a high projection | 3 |
3 | Antennal clava 7-segmented, A8–A12 strongly transverse; notauli weakly convergent anteriorly or subparallel, developed in posterior fourth or absent | S. crassiclavis |
– | Antennal clava 8-segmented, A8–A12 subquadrate or elongate; notauli distinctly divergent anteriorly and always developed at least in posterior third | S. formosus |
4 | (1) All femora broad, with very short stalks (fig. 5, 9; fig. 11, 9; fig. 14, 2 in |
5 |
– | All femora slender, with long stalks (fig. 8, 1, 8 in |
8 |
5 | Antenna with abrupt 6-segmented clava, A3–A7 yellowish, A8–A13 dark brown (fig. 5, 4, 5 in |
S. compressus |
– | Antenna with non-abrupt clava, uniformly reddish-brown to black (fig. 11, 5; fig. 14, 4 and fig. 15, 3 in |
6 |
6 | Clava slender, A11 about 1.5 times as wide as A4 in dorsal view and about 1.25 times, in lateral view (fig. 14, 4 in |
S. nigriclavis |
– | Clava wider, A11 about twice as wide as A4 in dorsal view and about 1.75 times, in lateral view (fig. 11, 5 and fig. 15, 3, 4 in |
7 |
7 | Neck of prothorax with short longitudinal grooves posteriorly; notauli developed at least in posterior third of mesoscutum (fig. 15, 6 in |
S. rufitarsis |
– | Neck of prothorax entirely smooth (fig. 11, 3 in |
S. latus |
8 | (4). Base of T2 pubescent (fig. 3, 3 and fig. 12, 4 in |
9 |
– | Base of T2 bare (fig. 1, 4 in |
11 |
9 | Micropterous (fig. 1 C. fig. 3 in |
S. antennatus |
– | Macropterous (Fig. |
10 |
10 | Propodeum with deep emargination between plicae, plicae slightly convergent posteriorly (Fig. |
S. brevimalaris sp. nov. |
– | Propodeum with not deep emargination between plicae, plicae not convergent posteriorly (Fig. |
S. flavecorpus sp. nov. |
11 | (8). T2 with numerous scattered long setae (fig. 6, 1 in |
S. cursor |
– | T2 bare; all ocelli present | 12 |
12 | Propodeum with deep arcuate emargination of posterior margin between plicae in dorsal view (fig. 2, 2; fig. 9, 6; fig. 10, 8 and fig. 17, 3 in |
13 |
– | Propodeum with weak arcuate emargination of posterior margin between plicae in dorsal view (fig. 1, 4; fig. 7, 1 and fig. 13, 4 in |
19 |
13 | Sternaulus complete (fig. 17, 1 in |
14 |
– | Sternaulus absent at least medially (fig. 9, 7 in |
17 |
14 | Head in front view with transverse wrinkles on antennal shelf (fig. 8, 2 in |
S. flavipes |
– | Head in front view without wrinkles on antennal shelf (fig. 2, 1 and fig. 17, 4 in |
15 |
15 | А3–А6 pale brown and clava black; tentorial pit absent to very tiny (punctiform) (fig. 2, 1 in |
S. annulicornis |
– | А1–А13 black; tentorial pit distinct (Fig. |
S. stigmaticalis |
16 | (13). Notauli usually absent, when rarely present, then expressed only in the form of two narrow incisions; malar sulcus totally absent | S. integer |
– | Notauli present in the form of broad grooves posteriorly; malar sulcus present, partly developed or fully visible in the form of a shallow groove | 17 |
17 | Neck of prothorax pubescent anteriorly (fig. 4, 7 in |
S. bipunctatus |
– | Neck of prothorax bare anteriorly (fig. 9, 6 in |
18 |
18 | Female antennae distinctly bicolor with abrupt 5-segmented clava (Fig. |
S. hemipterus |
– | Female antennae more or less monochrome with non-abrupt clava (Fig. |
S. thomsoni |
19 | (12). Notauli in the form of short grooves on mesoscutum posteriorly (fig. 1, 4 in |
S. abnormis |
– | Notauli totally absent; malar sulcus absent or incompletely developed (fig. 7, 2, 5 and fig. 13, 2 in |
20 |
20 | Head in dorsal view with temples parallel behind eyes (fig. 13, 5 in |
S. modestus |
– | Head in dorsal view with temples receding behind eyes (fig. 7, 8 in |
21 |
21 | Front behind scapus with two small oval and not deep holes (as in Fig. |
S. diversus |
– | Front behind scapus smooth (as in Fig. |
S. politus sp. nov. |
Female Spilomicrus stigmaticalis (ZSM-HYM-42423-H02, BOLD:ADS1706). A lateral; B dorsal; C face; D scutellum highlighted red, arrows mark the basal broadening; E antenna; F wing.
Female Spilomicrus hemipterus (ZSM-HYM-42322-F02; BOLD:ADM6694). A lateral; B dorsal; C face; D antenna.
Female Spilomicrus thomsoni (ZSM-HYM-42321-H08; BOLD:ADF4747). A lateral; B dorsal; C face; D antenna.
Males (males of S. cursor and S. nigriclavis unknown) |
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1 | Ventral margin of clypeus with pointed or rounded deflexed median projection; mandibles short, with upper tooth much shorter than lower tooth | 2 |
– | Ventral margin of clypeus with small rounded reflexed median projection (fig. 17, 2 [arrow], 4 in |
4 |
2 | In front view, ventral margin of clypeus rounded, blunt; A4 with moderately deep, curved emargination; mesosoma distinctly depressed, no more than 0.8 times as high as wide, mesoscutum weakly convex; median propodeal keel low, hardly raised anteriorly | S. sanbornei |
– | In front view, ventral margin of clypeus triangular, acuminate; A4 with at most a shallow emargination; mesosoma less depressed, at least 0.9 times as high as wide, mesoscutum strongly convex; median propodeal keel raised anteriorly to form a high projection | 3 |
3 | Eye sparsely hairy; А4 with carina over-reaching 0.7 of the segment | S. crassiclavis |
– | Eye bare; А4 with carina not over-reaching basal half of the segment | S. formosus |
4 | (1). Clypeus transverse, more than twice as wide as high (fig. 5, 2; fig. 11, 1 and fig. 15, 2 in |
5 |
– | Clypeus rounded, less than twice as wide as high (fig. 1, 2; fig. 2, 1 and fig. 4, 3 in |
8 |
5 | A4 distinctly longer than A3 | S. stigmaticalis |
– | A4 distinctly shorter than A3 | 6 |
6 | A5–A12 at least twice as long as wide (fig. 5, 3 in |
S. compressus |
– | A5–A12 at most 1.5 times as long as wide (fig. 11, 6 and fig. 15, 5 in |
7 |
7 | Neck with short longitudinal grooves posteriorly; notauli developed at least in posterior half of mesoscutum (fig. 15, 6 in |
S. rufitarsis |
– | Neck entirely smooth (fig. 11, 3 in |
S. latus |
8 | (4). Base of T2 pubescent (fig. 3, 3; fig. 12, 4 in |
9 |
– | Base of T2 bare (fig. 1, 4 in |
12 |
9 | A4 without emargination and keel (fig. 16, 6 in |
S. antennatus |
– | A4 with emargination and keel (fig. 12, 8, 9 in |
10 |
10 | Malar space 0.2–0.22 times as long as largest diameter of eye (Fig. |
S. brevimalaris sp. nov. |
– | Malar space more than 0.42 times as long as largest diameter of eye (Fig. |
11 |
11 | Head narrower than mesosoma (Fig. |
S. lusitanicus |
– | Head as wide as to wider than mesosoma in dorsal view (Fig. |
S. flavecorpus sp. nov. |
12 | (8). A3 distinctly longer than А4 (fig. 4, 2; fig. 9, 8 and fig. 10, 6, 7 in |
13 |
– | А3 as long as or shorter than А4 (fig. 1, 7; fig. 2, 7; fig. 7, 4; fig. 12, 9 and fig. 17, 10 in |
16 |
13 | Notauli present (fig. 4, 1 and fig. 9, 6 in |
14 |
– | Notauli absent (fig. 10, 8 in |
S. integer |
14 | Neck of prothorax pubescent anteriorly (fig. 4, 7 in |
S. bipunctatus |
– | Neck of prothorax bare anteriorly (fig. 9, 5, 6 in |
15 |
15 | A4 0.65 times as long as A3, widened apically with keel reaching 0.7 of the segment length (Fig. |
S. hemipterus |
– | A4 0.73 times as long as A3, cylindrical with keel reaching 0.55 of the segment length (Fig. |
S. thomsoni |
16 | (12). А4 with projection at base of keel and with bare smooth area along this keel (fig. 2, 7; fig. 8, 6, 7 and fig.17, 10 in |
17 |
– | А4 without projection at base of keel and without bare smooth area along this keel (fig. 1, 7; fig. 7, 4 and fig. 13, 6 in |
19 |
17 | Propodeum with deep arcuate emargination of posterior margin between plicae in dorsal view (fig. 8, 8 and fig. 17, 3 in |
12 |
– | Propodeum with weak arcuate emargination of posterior margin between plicae in dorsal view; А4 in lateral view wider than А3 and А5; pubescence of А3–А13 more dense, recumbent (fig. 2, 7 in |
S. annulicornis |
18 | Head in front view with transverse wrinkles on the top of antennal shelf (fig. 8, 2); antenna pale brown to brown, emargination on А4 shallow (fig. 8, 6 in |
S. flavipes |
– | Head in front view without transverse wrinkles on the top of antennal shelf (fig. 17, 4 in |
S. stigmaticalis |
19 | (16). Notauli developed in the form of short posterior grooves (fig. 1, 4 in |
S. abnormis |
– | Notauli totally absent (fig. 7, 1 and fig. 13, 4 in |
20 |
20 | Head in dorsal view subrectangular, with temples parallel behind eyes (fig. 13, 5 in |
S. modestus |
– | Head in dorsal view with temples receding behind eyes (fig. 7, 8 in |
21 |
21 | Front behind scapus with two small oval and not deep holes (Fig. |
S. diversus |
– | Front behind scapus smooth (Fig. |
S. politus sp. nov. |
Male antennae.
Male Spilomicrus stigmaticalis (ZSM-HYM-42320-F08, BOLD:ADS1706). A lateral; B dorsal.
DNA barcoding is revolutionising taxonomy research, especially when researchers are dealing with hyper- and cryptic-diverse insect taxa of small body size and variable morphological characters (
Another questionable case we faced was Spilomicrus stigmaticalis Westwood. While only an insufficient difference could be examined between the two haplotypes of the female, the males could not be distinguished morphologically at all. Interspecific variation was detected to be relatively low, while the intraspecific variation was rather high. Incomplete lineage sorting might be a reason for that, since allopatric/geographic factors, as well as seasonality could be ruled out. Taking both factors, genetics and morphology, into account, we decided to keep those two BINs in one species, S. stigmaticalis.
On the other hand, S. thomsoni was a relatively clear case. The two BINs (BOLD:ADF4747, BOLD:ADX1651), corresponding with the morphological determination, were genetically close (0.1%), while the taxon could be separated from S. hemipterus both genetically and morphologically.
There are still many taxonomic questions remaining regarding the Palearctic species of Spilomicrus. The high level of the sexual dimorphism in S. crassiclavis (
We are grateful to Dr. Rune Bygebjerg (MZLU) and Dr. Hege Vårdal (NHRS) for the photos of the type specimens and to Dr. Zoltan Vas (HNHM) and Dr. Claire Villemant (MNHN) for their help and providing the opportunity to examine the type material.
Phylogenetic ML-tree of 45 Spilomicrus sequences with the outgroup Labolips innupta.
This table lists all the location data for each specimen that was caught within the project. Not listed are the lectotypes stored at other museums that were investigated. All the available information is printed on the labels in the image tables.