A new species of Pristaulacus Kieffer (Hymenoptera: Aulacidae) from Baltic amber

Insect Systematics & Evolution 40 (2009) 201–207

brill.nl/ise

A new species of Pristaulacus Kieffer (Hymenoptera: Aulacidae) from Baltic amber John T. Jenningsa,* and Lars Krogmannb a

Australian Centre for Evolutionary Biology and Biodiversity, and School of Earth and Environmental Sciences, The University of Adelaide, SA 5005, Australia b State Museum of Natural History Stuttgart, Entomology, Germany *Corresponding author, e-mail: [email protected]

Abstract Fossil species of Pristaulacus are uncommon, with just two known species, P. mandibularis Brues and P. praevolans Brues from Baltic amber, and three species, P. bradleyi (Brues), P. rohweri Brues and P. secundus (Cockerell), from the Florissant fossil beds, Colorado, USA. Here we provide a detailed description of Pristaulacus velteni sp.n., the third fossil species known from Baltic amber. Keywords Hymenoptera, Evanioidea, aulacid wasp, Eocene, Pristaulacus velteni sp.n.

Introduction The Evanioidea are generally considered to comprise three extant families, Aulacidae, Gasteruptiidae and Evaniidae (e.g., Naumann 1991, Gauld & Bolton 1996; Jennings et al. 2004a,c; Turrisi et al. 2009), although Townes (1950) and Zhang & Rasnitsyn (2008), for example, treat Aulacidae as a subfamily of Gasteruptiidae. As well, two extinct families, Praeaulacidae and Andreneliidae, have been placed in this superfamily (Rasnitsyn 1972, 2002; Rasnitsyn & Martínez-Delclòs 2000; Zhang & Rasnitsyn 2008), although Engel (2006) considers the latter family within Evaniidae. Other basal lineages of extinct evanioid wasps of uncertain affinity include Baissidae (including Manlaya Rasnitsyn 1980; Zhang and Rasnitsyn 2004), placed as sister to the remaining Aulacidae (Nel et al. 2004), or as basal sister-group of the clade Aulacidae + Gasteruptiidae (Engel 2006) or as a subfamily of Gasteruptiidae s.l. (Zhang & Rasnitsyn 2007), and the Kotujellidae (Rasnitsyn 1975), currently placed within Aulacidae (Rasnitsyn 1980). Aulacidae are currently divided into three extant genera, Aulacus, Panaulix and Pristaulacus, although recent phylogenetic analyses based on morphology (Turrisi et al. 2009) indicate that Aulacus is paraphyletic to the clade Pristaulacus + Panaulix. © Koninklijke Brill NV, Leiden, 2009

DOI 10.1163/187631209X440069

202

J.T. Jennings and L. Krogmann / Insect Systematics & Evolution 40 (2009) 201–207

Aulacidae are also known from fossils, with some 36 described species (Basibuyuk et al. 2002; Nel et al. 2004; Jennings et al. 2004b; Zhang & Rasnitsyn 2007). Approximately half of these species are from the Cenozoic, with records from the Upper Eocene of the Isle of Wight and Baltic amber, Lower Eocene Paris basin amber, and from the Oligocene of North America (Nel et al. 2004). As well, Hyptiogastrites electrinus Cockerell, from the Lower Cretaceous of Myanmar, is possibly a sister taxon to Aulacidae (Jennings et al. 2004b), or represents a distinct ancestral lineage of Aulacidae, the Hyptiogastritinae (Engel 2006). This group is characterized by a distinct occipital carina and hind coxal ovipositor guide, and the absence of fore wing veins 1rs-m, 2rs-m, 2m-cu and a (Jennings et al. 2004b; Engel 2006). Fossil species of Pristaulacus are rare, with just five species known, each from a single specimen: P. mandibularis (Brues 1933) and P. praevolans (Brues 1923) from Baltic amber, and P. bradleyi (Brues 1910), P. rohweri (Brues 1910) and P. secundus (Cockerell 1916) from the Florissant fossil beds, Colorado, USA. Here we describe a new fossil species of Pristaulacus, being just the third species known from Baltic amber. Material and Methods The specimen was examined under a Leica Z16 APO Microscope and images taken with a digital camera (Leica DFC 490) attached to it. Images were processed with Automontage (Synoptics). Terms for general morphology follow Jennings and Austin (1994), and that for wing venation follows the modified Comstock–Needham system after Sharkey (1988), but with some modifications, and using the nomenclature of van Achterberg (1979) for cells. Systematic Palaeontology Family Aulacidae Hedicke, 1939 The systematic history of Aulacidae is reviewed in Smith (2001). Genus Pristaulacus Kieffer, 1900 For a complete list of generic synonomies, see Turrisi (2007). Pristaulacus velteni sp.n. (Figs 1, 2A-E) Diagnosis Wasp of medium size; 9.1 mm in length, excluding ovipositor; fore wing 6.15 mm in length; occipital carina well-defined; antenna with 12 flagellomeres; hind claw pectinate, with 3 medial tooth-like processes (basal process not visible) (Fig. 2E); fore wing vein 2-Rs+M short compared with 1-Rs+M, marginal cell 3.8 times as long as wide, vein 2r-m absent, 3r-m present (Fig. 2D). Occurrence Eocene Baltic amber. Holotype female deposited in Staatliches Museum für Naturkunde Stuttgart, Germany (SMNS, Coll. No. BB-2393).

J.T. Jennings and L. Krogmann / Insect Systematics & Evolution 40 (2009) 201–207

203

Fig. 1. Pristaulacus velteni sp.n. female holotype specimen in lateral view. This figure is published in colour in the online edition that can be accessed via http://www.brill.nl/ise

Description Female. Head 1.4× longer than wide when viewed laterally (Figs 1, 2B); face and vertex with numerous small piliferous punctures; posterior margin of head slightly concave in dorsal view; occipital carina present, narrow, less than 0.2× diameter of lateral ocellus; malar space 0.36× height eye; clypeus 3.1× as wide as high, margin sinuate, without distinct medial process; distance from lateral ocellus to eye margin 1.0× distance between lateral ocelli; antenna with 12 flagellomeres; scape 1.5× length pedicel; first flagellomere 1.7× as long as scape, 0.7× as long as second flagellomere (Fig. 2A). Mesoscutum in lateral view angular antero-dorsally (Fig. 2B, C), medial and lateral lobes coarsely transverse-carinate, admedial lines absent; scutellum coarsely transversecarinate; axilla narrow, single carina on posterior margin; mesepimeron broad, smooth; parascutal carina expanded, with tooth-like lateral projection; propodeum in posterior view, with two complete sub-medial carinae and incomplete median carina; subapical transverse hind coxal groove on inner surface indistinct; hind trochanter with dorsal groove; prefemur on hind leg present; hind femur 0.8× length hind tibia; hind tarsomeres 1–4 with ventro-apical pecten of short robust spines, tarsomere 1, 3.0× length tarsomere 2; tarsomere 2, 1.4× length tarsomere 3; tarsomere 3, 2.2× length tarsomere 4; tarsomere 4, 0.7× length tarsomere 5; hind tarsal claw 0.4× length tarsomere 5; hind claw pectinate, with 3 medial tooth-like processes plus long apical tooth-like process,

204

J.T. Jennings and L. Krogmann / Insect Systematics & Evolution 40 (2009) 201–207

Fig. 2. Pristaulacus velteni sp.n. female holotype specimen. (A) Head in frontal view, (B) head and anterior mesosoma in lateral view, (C) mesosoma in lateral view, (D) left forewing in ventral view (dotted line indicates ‘normal’ position of reduced 2r-m) and (E) right mesotarsus in lateral view. Abbreviations: at, apical tooth-like process; mt, median tooth-like process; ped, pedicel; F1, first flagellomere; F2, second flagellomere; msc, mesoscutum; sc, scape; tarn, n-th tarsomere. This figure is published in colour in the online edition that can be accessed via http://www.brill.nl/ise

basal process not visible (Fig. 2E); fore wing with vein 2r-m absent, but 3r-m present, vein 2-Rs+M short compared with 1-Rs+M, marginal cell 3.8× as long as wide (Fig. 2D); hind wing venation complete, R+Rs, M+Cu, Cu, r-m and 2-M present. Metasoma ovate in lateral view, 1.3× length of mesosoma; petiole stocky, T1 and T2 smooth dorsally; ovipositor exserted but incomplete. Male unknown. Remarks This new species clearly belongs to Pristaulacus. The presence of pectinate hind claws is a characteristic common to all extant Pristaulacus species (Turrisi et al. 2009), although the number, shape and placement of the medial tooth-like processes vary. On the other hand, all extant Aulacus species have non-pectinate hind claws (see, for example,

J.T. Jennings and L. Krogmann / Insect Systematics & Evolution 40 (2009) 201–207

205

Jennings et al. 2004a). Also, the presence of a complete occipital carina is common to the majority of Pristaulacus, although the medial part is obsolete in some Australian and Afrotropical species (see Jennings et al. 2004a; Turrisi 2006), but it is completely absent in Aulacus. Pristaulacus bradleyi, P. rohweri and P. secundus are known from the Florissant fossil beds which are from the latest Eocene to lowermost Oligocene (approx. 35 million years ago) (Evanoff et al. 2001), whereas the other two fossil Pristaulacus species occur in Middle Eocene Baltic amber (approx. 38–54 million years ago) (KosmowskaCeranowicz 1987), although the age of Baltic amber may be Late Eocene (Perkovsky et al. 2007). Pristaulacus velteni is 9.1 mm in length, whereas P. bradleyi is a much larger species, estimated by Brues (1910) to be 18 mm in length (the metasoma is missing). Pristaulacus rohweri is a smaller species and is 7 mm in length. The length of P. secundus cannot be estimated as the specimen is incomplete. However, it has smaller wings than P. bradleyi (Cockerell 1916), suggesting that it is probably comparable in size with P. velteni sp.n. Pristaulacus velteni sp.n. also differs from these three species in wing venation. As indicated by Brues (1910), P. bradleyi has a quite different fore wing venation from other Pristaulacus species, particularly the unusual shape of the first submarginal cell caused by the lack of vein 2-Rs+M (see Brues 1910: Fig. 19). In P. velteni sp.n., 2-Rs+M is present but distinctly shortened (Fig. 2D), and the marginal cell is wider than in any other fossil Pristaulacus, being 3.8 times as long as wide compared with: 3.1 times in P. bradleyi, 3.6 in P. secundus and 2.25 in P. rohweri. The fore wing of P. secundus is generally similar to that of P. velteni, sp.n., but vein 2r-m is present (Cockerell 1916: Fig. 9a), unlike P. velteni sp.n. which lacks 2r-m (Fig. 2D). Pristaulacus bradleyi and P. rohweri also lack vein 2r-m. Compared with the two Baltic amber species, P. velteni sp.n. is slightly shorter in length than both P. mandibularis and P. praevolans, which are 10 mm and 12 mm long respectively (Brues 1923, 1933). This new species differs from P. mandibularis in a number of character states including: anterior margin of clypeus produced into a median tooth in P. mandibularis (Brues 1933) which is absent in P. velteni sp.n. (Fig. 2A), mesonotum transverse-carinate anteriorly on sides but not obviously carinate medially in P. mandibularis (Brues 1933), whereas it is distinctly carinate with coarse ridges in P. velteni sp.n. and scutellum coarsely transverse-carinate in P. velteni sp.n., but finely reticulate in P. mandibularis (Brues 1933). Unfortunately the hind claws are missing on P. mandibularis (Brues 1933), so no comparison can be made between the species. Pristaulacus velteni sp.n. differs from P. praevolans in a number of other character states including: P. praevolans has two medial tooth-like processes and a weak basal process on the hind claw (Brues 1923), whereas P. velteni sp.n. has three medial processes (Fig. 2E), and the metasoma of P. praevolans is about same length as the mesosoma (Brues 1923), whereas it is 1.3× as long in P. velteni sp.n. This species is named after Jürgen Velten (Idstein, Germany) who kindly donated the amber specimen to the collection of SMNS.

206

J.T. Jennings and L. Krogmann / Insect Systematics & Evolution 40 (2009) 201–207

Acknowledgements We are grateful to Jürgen Velten who brought the specimen to our attention and donated the holotype to the amber collection of SMNS. Karin Wolf-Schwenninger (SMNS) kindly assisted us with preparation of the specimen. We also thank G.F. Turrisi and A.P. Rasnitsyn for their valuable comments on the manuscript.

References Basibuyuk, H.H., Rasnitsyn, A.P., Fitton, M.G. & Quicke, D.L.J. (2002) The limits of the family Evaniidae (Insecta: Hymenoptera) and a new genus from Lebanese amber. Insect Systematics & Evolution 33: 23–34. Brues, C.T. (1910) The parasitic Hymenoptera of the Tertiary of Florissant, Colorado. Bulletin of the Museum of Comparative Zoology at Harvard College 54: 1–125+1 plate. Brues, C.T. (1923) Some new fossil parasitic Hymenoptera from Baltic amber. Proceedings of the American Academy of Arts and Sciences 58: 327–346. Brues, C.T. (1933) The parasitic Hymenoptera of the Baltic amber. Part I. Bernstein-Forschungen 3: 4–178. Cockerell, T.D.A. (1916) Some American fossil insects. Proceedings of the U.S. National Museum 51: 89–107. Deans, A.R., Basibuyuk, H.H., Azar, D. & Nel. A. (2004) Descriptions of two new Early Cretaceous (Hauterivian) ensign wasp genera (Hymenoptera: Evaniidae) from Lebanese amber. Cretaceous Research 25: 509–516. Engel, M.S. (2006) Two ensign wasps in Cretaceous amber from New Jersey and Myanmar (Hymenoptera: Evaniidae). Polskie Pismo Entomologiczne 75: 443–454. Evanoff, E., McIntosh, W.C. & Murphey, P.C. (2001) Stratigraphic summary and 40Ar/39Ar geochronology of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature and Science 4: 1–16. Gauld, I.D. & Bolton, B. (1996) The Hymenoptera. 2nd edn. Oxford University Press, Oxford. Jennings, J.T. & Austin, A.D. (1994) Revision of the genus Crassifoenus Crosskey (Hymenoptera: Gasteruptiidae: Hyptiogastrinae), with a description of a new species from Western Australia. Records of the Western Australian Museum 16: 575–591. Jennings, J.T., Austin, A.D. & Stevens, N.B. (2004a) The aulacid wasp fauna of Western Australia with descriptions of six new species. Records of the Western Australian Museum 22: 115–128. Jennings, J.T., Austin, A.D. & Stevens, N.B. (2004b) Hyptiogastrites electrinus Cockerell, 1917, from Burmese amber: redescription and its placement within the Evanioidea (Hymenoptera). Journal of Systematic Palaeontology 2: 127–132. Jennings, J.T., Austin, A.D. & Stevens, N.B. (2004c) First record of Aulacidae (Hymenoptera: Evanioidea) from New Caledonia with descriptions of three new species of Aulacus Jurine. Journal of Australian Entomology 43: 346–352. Kosmowska-Ceranowicz, B. (1987) Charakterystyka mineralogiczno- petrograficzna osadów bursztynonośnych Eocenu w rejonie Chłapowa oraz osadów Paleogenu Polnocnej Polski. Biuletyn Instytutu Geologicznego 356: 29–50. [in Polish] Naumann, I.D. (1991) Hymenoptera (Wasps, bees, ants, sawflies). In: Naumann I.D. (Ed.) The Insects of Australia, Volume 2. CSIRO, Melbourne University Press, Carlton, Vic, pp. 916–1000. Nel, A., Waller, A. & De Ploëg, G. (2004) An aulacid wasp in the Lowermost Eocene amber from the Paris Basin (Hymenoptera: Aulacidae). Geologica Acta 2: 67–74. Perkovsky, E.E., Rasnitsyn, A.P., Vlaskin, A.P. & Taraschuk, M.V. (2007) A comparative analysis of the Baltic and Rovno amber arthropod faunas: representative samples. African Invertebrates 47: 229–245.

J.T. Jennings and L. Krogmann / Insect Systematics & Evolution 40 (2009) 201–207

207

Rasnitsyn, A.P. (1972) Praeaulacidae (Hymenoptera) from the Upper Jurassic of Karatau. Paleontologicheskii Zhurnal 1: 72–87. [in Russian] Rasnitsyn, A.P. (1975) Hymenoptera Apocrita from the Mesozoic, Trudy Paleontologicheskogo Instituta Akademii nauk SSSR. Nauka, Moscow. Rasnitsyn, A.P. (1980) Contribution to the knowledge of the system of the family Aulacidae (Hymenoptera) in connexion with a new find in Neocom of Manalay. Trudy Sovmestnaya Sovetsko-Mongol’skaya paleontologischeskaya Expeditsiya, Moscow 13: 65–67. [in Russian] Rasnitsyn, A.P. (2002) Suborder Vespidea Laicharting, 1781. In: Rasnitsyn, A.P. & Quicke, D.L.J. (Eds), History of Insects. Kluwer, Dordrecht, pp. 242–254. Rasnitsyn, A.P. & Martínez-Delclòs, X. (2000) Wasps (Insecta: Vespida = Hymenoptera) from the Early Cretaceous of Spain. Acta Geologica Hispanica 35: 65–95. Sharkey, M. (1988) Ichneumonoid wing venation. Ichnews 11: 2–12. Smith, D.R. (2001) World catalog of the family Aulacidae (Hymenoptera). Contributions on Entomology, International 4: 261–319. Townes, H.K. (1950) The Nearctic species of Gasteruptiidae (Hymenoptera). Proceedings of the United States National Museum 100: 85–145. Turrisi, G.F. (2006) Revision of the Afrotropical species of Pristaulacus Kieffer, 1900 (Hymenoptera: Aulacidae). Insect Systematics & Evolution 37: 27–38. Turrisi, G.F. (2007) Revision of the Palaearctic species of Pristaulacus Kieffer, 1900 (Hymenoptera: Aulacidae). Zootaxa 1433: 1–76. Turrisi, G.F., Jennings, J.T. & Vilhelmsen, L. (2009) Phylogeny and generic concepts in the parasitoid wasp family Aulacidae (Hymenoptera: Evanioidea). Invertebrate Systematics, 23: 27–59. van Achterberg, C. (1979) A revision of the Subfamily Zelinae auct. (Hymenoptera, Braconidae). Tijdschrift voor Entomologie 122: 241–479. Zhang, H. & Rasnitsyn, A.P. (2007) Nevaniinae subfam. n., a new fossil taxon (Insecta: Hymenoptera: Evanioidea: Praeaulacidae) from the Middle Jurassic of Daohugou in Inner Mongolia, China. Insect Systematics & Evolution 38: 149–166. Zhang, H. & Rasnitsyn, A.P. (2008) Middle Jurassic Preaulacidae (Insecta; Hymenoptera: Evanioidea) of Inner Mongolia and Kazakhstan. Journal of Systematic Palaeontology 6: 1–25.