Revision of the south Asian amisegine genus Cladobethylus Kieffer, 1922 (Hymenoptera, Chrysididae, Amiseginae)

The south Asian genus Cladobethylus Kieffer, 1922, is reviewed, with a key to the species. Five new species are described, one from the island of Borneo ( Cl. darlingi ), two from Thailand ( Cl. densepunctatus and Cl. thailandicus ), one from Korea ( Cl. koreensis ), and one from Sumatra ( acehensis ).


Revision of the south Asian amisegine
genus Cladobethylus Kieffer, 1922 (Hymenoptera, Chrysididae, Amiseginae) Introduction Cladobethylus Kieffer, 1922, is one of the most widespread and speciose of the south Asian amisegine genera, with 11 species, including the ones newly described below.There are undoubtedly more species to be discovered and it would not be surprising to find species in southern China, Cambodia and other islands in Indonesia, such as Sulawesi.The current distribution of the genus is probably more a reflection of where collecting has been done and the techniques used than the actual distribution.Based on personal observation, as with most other amisegines, males are more commonly collected, particularly in Malaise traps and by sweep netting.Females are rarer in collections, and are more likely to be collected in bowl and pitfall traps.This may have more to do with the environments the two sexes typically inhabit than any actual differences in numbers.Females may spend most of their time searching in and on leaf litter for walking stick eggs, whereas males fly and sit on vegetation above the ground searching for females, and are more likely to be caught in a Malaise trap.
The majority of Cladobethylus species are moderate-sized amisegine wasps ranging from 2-4 mm long.However, one species, Cl. insularis Kimsey & Dewhurst, from Papua New Guinea is much larger, ranging from 5-7 mm in length.This is also the only species, where a host is known.It parasitizes the eggs of Eurycantha insularis Lucas (Phasmatodea: Phasmatidae) (Kimsey et al. 2012).

Materials and methods
Specimens were borrowed from and/or types are deposited in the following institutions, which are given in the descriptions below using their acronyms: AEI -American Entomological Institute, Logan, Utah, USA; BME -Bohart Museum of Entomology, University of California, Davis, USA; BMNH -The Natural History Museum, London, U.K.; BPBM -B.P. Bishop Museum, Honolulu, Hawaii, USA; MZB -Museum Zoologi Bogor, Chibinong, Indonesia; QSBG -Queen Sirikit Botanical Garden, Chiang Mai, Thailand; ROM -Royal Ontario Museum, Toronto, Ontario, Canada; USNM -U. S. National Museum of Natural History, Washington, D. C., USA.
Morphological terminology follows that used by Kimsey and Bohart (1991).The acronym MOD is used for midocellus diameter, which is the width of the midocellus from side to side in front view.The interocular distance is measured across the narrowest part of the face in front view, using the greatest width of the eye in front view for comparison.Antennal articles are measured at the point of greatest breadth and compared with the total length of the article.Postocular distance is measured across the greatest width of the head laterally behind the eye margin in dorsal view.
Diagnosis.The most distinctive feature of the genus Cladobethylus is the impunctate welt that extends along the midline of the vertex from near the midocellus to the occipital carina.This feature is only shared with species of Magdalium Kimsey and Senesega Kimsey, although it is less well developed in these genera.Cladobethylus can be distinguished from those genera by the lack of an omaulus on the mesopleuron and the usually biangulate genal area, which is evenly curved in Magdalium and Senesega.
In addition, Cladobethylus species have at most a weakly defined scrobal sulcus.The scrobal sulcus in Magdalium and Senesega is well defined.Description.Vertex with low impunctate welt along midline from midocellus to occiput; eye with short sparse setulae; scapal basin usually with at least some crossridging; malar space with vertical sulcus extending from ventral eye margin to mandibular articulation; occipital carina present; posterior margin of head behind eye usually biangulate in side view; male flagellum elongate and cylindrical; female flagellum short, fusiform and usually flattened on one surface; pronotum with narrow posteromedial groove; mesopleuron without scrobal sulcus or omaulus; scutum with notauli and parapsides; both sexes fully winged, forewing Rs extended by evenly curved dark streak, costal cell widest medially, stigma large and broad, R 1 not indicated, medial vein arising before cu-a; propodeum with long dorsal surface and abrupt posterior declivity, lateral propodeal angle undeveloped; hindcoxa with dorsobasal carina; tarsal claws edentate in females, with one small subbasal tooth and subbasal angle in males.
Etymology.The species is named after Aceh Province on the island of Sumatra, Indonesia, where it was collected.
Diagnosis.Cladobethylus aquilus males most closely resemble those of Cl. ceylonicus and Cl.insularis based on flagellomere I 4.0-4.5×as long as broad, the dark brown antenna, nearly impunctate metasomal tergum II, and reduced cross ridging in the scapal basin.However, Cl. aquilus can be distinguished from those and other species by the combination of the longer subantennal distance, lack of metallic blue highlights, slightly medially lobate apical flagellomeres, and flagellomere XI 6× as long as broad.Females most closely resemble those of Cl. insularis and Cl.thailandicus based on the bicolored antenna and flagellomere II about as long as broad.They can be distinguished from these and other species by the scapal basin lacking cross ridges, bicolored legs and densely punctate metasomal tergum II.
Diagnosis.The most distinctive features of Cl. ceylonicus are the long first and second flagellomeres, a characteristic shared with Cl. acehensis and Cl.densepunctatus.Howev-er, Cl. ceylonicus can be distinguished from these and other species by the combination of these features, along with the densely punctate second metasomal tergum, asymmetrical apical male flagellomeres and short subantennal distance.Male description.Body (Figs 8,9).Length 4.0 mm.Head.Scapal basin impunctate, with short strip of cross-ridges on either side of narrow medial stripe; clypeus short and subtruncate apically, subantennal distance 0.4 MOD long; malar space 3.8 MOD long wide; head about as long as wide; interocular distance 1.9× eye width in front view; midocellus 2.2 MOD from ocular margin; ocelli arranged in isosceles triangle; hindocelli separated from ocular margin by 1 diameter; postocular distance 1.6 MOD wide; flagellomere I 4.5× as long as broad, setae 0.5× as long as flagellomere breadth; flagellomere II 3.3× as long as broad; flagellomere XI 5× as long as broad; flagellomeres VIII-X asymmetrically bulging submedially.Mesosoma.Pronotum about 1.1× as long as scutum; mesopleural punctures contiguous; metapleuron dorsally cross-ridged, ventrally polished, impunctate; propodeum laterally cross-ridged dorsally, ventrally polished, impunctate, with subrectangular posteromedial enclosure.Metasoma.Tergum I with few, widely scattered tiny punctures; tergum II with large patch of punctures 1-2 puncture diameters apart, separated by medial longitudinal stripe; terga III-V with dense, tiny, nearly contiguous punctures, with impunctate apical margin.Color.Body black, with metallic blue highlights; legs yellow; antenna orange; wing membrane brown-tinted, veins dark brown.

Cladobethylus cruciger
Female description (based on Kieffer's description of Cl. coeruleus).Body length 4.0 mm.Head.Flagellomere I 2.5× as long as broad; flagellomere II 1.5× as long as broad; flagellomere X more than twice as long as broad.Metasoma.Third tergum closely and rather finely punctured.Color.Body black, except mesosomal dorsum with metallic blue highlights; legs including coxae yellow.
Discussion.The description above is based on Kieffer's descriptions of Cl. coeruleus and Cl.cruciger, plus a single male specimen from the island of Mindanao that fits Kieffer's description.
Kieffer's Cladobethylus coeruleus is synonymized here with Cl. cruciger as his description of coeruleus matches the shared characteristics between male and female Cladobethylus seen in other species where both sexes are known.Both specimens are from Mindanao.In the female description, Kieffer refers to punctation of the third tergum, but it is not clear whether he counted the propodeum as the first tergum.
The subspecies Cl. antennalis might be a valid species based on the differences in flagellomere dimensions and collection location on a separate island.In Kieffer's description flagellomere I is 3.5× along as broad and flagellomere II 3× as long as broad in Cl. cruciger s. s., but in Cl. antennalis flagellomere I is 3× as long as broad and II twice as long as broad.However, without access to the types it is impossible to be certain.Cladobethylus by the combination of the malar space 3.5 midocellar diameters wide or wider, interocular distance more than twice eye width in front view, flagellomere I less than 3.5× as long as broad, flagellomere II less than 2.5× as long as broad, and flagellomere XI more than 6.5× as long as broad.
Etymology.The species is named for the unusually dense, coarse punctation on the head and thorax.
Diagnosis.Cladobethylus gilbus shares the nearly impunctate second metasomal tergum with Cl. darlingi, Cl. acehensis and Cl.densepunctatus.In males it can be distinguished from those species by the narrow malar space (less than 2.8 midocellar diameters), flagellomere I less than 4× as long as broad, flagellomere II more than 2.5× as long as broad, flagellomere XI 7× as long as broad and postocular distance less than 1 MOD.Females have the basal three flagellar segments paler than the rest of the flagellum, very short flagellomeres I and II and malar space 4.5 midocellar diameters wide (shared with Cl. aquilus).Male description.Body (Figs 21,22).Length 3.5-4.0mm.Head.Scapal basin with narrow band of cross-ridges separated by smooth medial band; head venter without ovoid foveae along midline of gular area, gular area punctate; malar space 2.6 MOD long; head as wide as long; interocular distance 1.5× eye width in front view; midocellus 1.4 MOD from ocular margin; ocelli arranged in a nearly equilateral triangle; hindocelli separated from ocular margin by 1 diameter; subantennal distance 0.6 MOD; clypeal apex truncate to slightly concave; flagellomere I 3.6× as long as broad, setae length 0.4× flagellomere breadth; flagellomere II 2.6× as long as broad; flagellomere IX 7× as long as broad; flagellomeres VIII-X slightly lobate submedially.Mesosoma.Pronotum 0.9× as long as scutum; mesopleural punctures 0.2-0.5 puncture diameters apart; metapleuron with zone of cross ridging below hindwing base; propodeum laterally cross-ridged, posteromedially with subrectangular enclosure.Metasoma: terga I and II impunctate; terga III-V with basal band of dense, minute, contiguous punctures.Color: body black, except pronotum, scutum, scutellum and metanotal triangle with metallic blue highlights; antenna and legs including coxae yellow; wing membrane light brown tinted, with brown veins.
Distribution.MALAYSIA: Negeri Sembilan, Pasoh Forest Res.; 106 males and 50 females were examined (AEI, BME).Males and females were found in nearly every month over a two-year period of Malaise trap collecting at this site (Fig. 20).
Diagnosis.This species has several unusual diagnostic female features.The eye has very long dense setulae, the hindocelli are very close to the ocular margins and the malar space is very long.In addition, the pronotum is longer than the scutum, and the metapleuron is cross-ridged from wing base to coxa.
Male.Unknown.Female description.Body (Figs 30,31).Length: 2.5 mm.Head.Scapal basin smooth with short strip of cross-ridges on either side of broad medial stripe; clypeus short, broadly rounded; subantennal distance 0.6 MOD; malar space 5 MOD long; interocular distance 1.5-1.7×eye width in front view; face about as broad as long in front view; ocular setulae 1 MOD long or longer; midocellus 2.6 MOD from ocular margin; hindocellus separated from ocular margin by one diameter; ocelli arranged in nearly equilateral triangle; hindocelli separated from ocular margin by 1 hindocellar diameter; flagellomere I 2.3× as long as broad; flagellomere II 1.4× as long as broad.Mesosoma.Mesopleural punctures larger than on pronotum; metapleuron cross-ridged from hindwing base nearly to midcoxa; propodeum with posteromedial enclosure rough, subrectangular.Metasoma: terga I-V with tiny scattered punctures at least 4-6 PD apart laterally, nearly impunctate medially.Color: body black with bluish tints on vertex and pronotum; legs including coxae yellow; scape dark brown becoming paler distally; pedicel and flagellomeres I-III yellow, remaining flagellomeres brown; mandibles brown.
Distribution.JAPAN: Kyoto; 1 female (the holotype) examined (AEI).Diagnosis.This species has several unusual female features.The eye has very long dense setulae, the hindocelli are very close to the ocular margins and the malar space is very long.In addition, the pronotum is longer than the scutum, and the metapleuron is cross-ridged from wing base to coxa.Male.Unknown.Female description.Body (Figs 32,33).Length.2.5-3.0 mm.Head.Scapal basin smooth, polished with narrow strip of cross-ridges on either side of broad medial stripe; clypeus short, broadly convex apicomedially; subantennal distance 0.5 MOD wide; malar space 5 MOD long; interocular distance 1.2× eye width in front view; face about 3× as long as broad across midocellus in front view; ocular setulae 1.5 MOD long or longer; midocellus 3 MOD from ocular margin; hindocellus separated from ocular margin by 1 hindocellar diameter; ocelli arranged in isosceles triangle; postocular distance 0.5 MOD; flagellomere I 2.3× as long as broad; flagellomere II as long as broad; flagellomere X 1.5× as long as broad.Mesosoma.Mesopleural punctures larger than on pronotum, 0.2-1.0puncture diameters apart; metapleuron impunctate medially, transversely ridged, with smooth area medially; propodeum laterally impunctate and irregularly wrinkled, posteromedial enclosure ovoid.Metasoma: terga I-V with tiny scattered punctures at least 10 PD apart laterally, nearly impunctate medially.Color: body black without bluish tints; legs including coxae yellow; scape dark brown becoming paler distally; pedicel and flagellomeres I-III white, remaining flagellomeres brown; mandibles brown; wing base white, rest of wing membrane and veins brown.
Etymology.The species is named for the country of collection.Diagnosis.This is a distinctive species.Males can be distinguished by flagellomere XI 5× as long as broad (shared with Cl. ceylonicus and Cl.densepunctatus), scapal basin without cross-ridging, subantennal distance more than 1 midocellar diameter long and body lacking metallic bluish highlights.Females have a bicolored antenna, with the basal segments whitish and the apical segments dark brown, the postocular distance is less than one midocellar diameter, and the gular bridge lacks occipital foveae.Male description.Body (Figs 34,35).Length 2.5-4.0 mm.Head.Frons punctures 1-2 puncture diameters apart; scapal basin primarily smooth with short strip of cross-ridges on either side of broad medial stripe; clypeus long and truncate or slightly emarginate apically; subantennal distance 1.2 midocellar diameters; malar space 3.5 midocellar diameters long; head about as long as wide; midocellus 1.7 diameters from ocular margin; ocelli arranged in nearly equilateral triangle; hindocelli separated from ocular margin by one diameter; postocular distance 2.5 midocellar diameters; flagellomere I 4× as long as broad, setae 0.5× as long as flagellomere breadth; flagellomere II 3.2× as long as broad; flagellomere IX 5× as long as broad.Mesosoma.Pronotum about 0.9× as long as scutum; mesopleural punctures 0.5-1.0puncture diameters apart; metapleuron smooth, impunctate; propodeum laterally wrinkled, with posteromedial enclosure subrectangular.Metasoma.Tergum I essentially impunctate; tergum II with few, widely scattered, tiny punctures; terga III-V with dense, tiny punctures 0-0.5 puncture diameters apart, with impunctate apical margin.Color.Body black, except pronotum, scutum, scutellum and medial triangle of metanotum with metallic blue highlights; legs including coxae yellow to orange; antenna brown; wing membrane brown, veins dark brown.

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This is the largest bodied species of Cladobethylus, with males averaging about 1.5× the length of the known other species.Male Cl. insularis can be distinguished from Cl. aquilus, the only other species known from New Guinea by the pronotum without blue tints, a much broader zone of cross-ridging in the scapal basin, legs brown instead of yellow (Cl.aquilus) flagellomere I shorter (4× as long as broad versus 5× in Cl. aquilus), and flagellomere XI shorter (5× as long as broad versus 6× in Cl. aquilus).Features of female Cl.insularis not shared with other Cladobethylus include the bicolored antenna and legs, long clypeus (1 MOD long versus 0.3-0.6MOD in other species), and narrow distance between the midocellus and nearest eye margin (up to 2 MOD versus 2.6-2.7 MOD in the others).