A new family, Cretostylopidae Kathirithamby and Engel, is erected to accommodate the mid-Cretaceous species Cretostylops engeli Grimaldi and Kathirithamby. Based on this as well as various other recent works pertaining to the higher classification of Strepsiptera, a revised key is provide to the living and fossil families

Engel, MS

Kathirithamby, Jeyaraney

Journal of the Kansas Entomological Society

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A Revised Key to the Living and Fossil Families of Strepsiptera, with theDescription of a New Family, CretostylopidaeJEYARANEY KATHIRITHAMBY1 AND MICHAEL S. ENGEL2,3ABSTRACT: A new family, Cretostylopidae Kathirithamby and Engel, is erected toaccommodate the mid-Cretaceous species Cretostylops engeli Grimaldi and Kathirithamby.Based on this as well as various other recent works pertaining to the higher classification ofStrepsiptera, a revised key is provide to the living and fossil families.KEY WORDS: Strepsiptera, twisted-winged parasites, Cretostylops, fossil, identification keyRecently there has been increased interest in the higher classification of evolutionof the twisted-winged parasites (Strepsiptera). Indeed, many new changes have beenproposed by a variety of authors, particularly in regard to some new fossil andprimitive modern lineages (e.g., Grimaldi et al., 2005; Pohl and Beutel, 2005; Pohl etal., 2005; Bravo et al., 2009). Despite this fervor of activity, there has been no revisedmeans through which to identify this diversity. Accordingly, we provide a revised keyto the living and fossil families of Strepsiptera. The key is provided here to permitidentification not only of recently proposed families (Pohl and Beutel, 2005; Pohl etal., 2005; Bravo et al., 2009), but also a family newly established herein. It has beenlong known that the Cretaceous genus Cretostylops Grimaldi and Kathirithamby isrepresentative of a distinct group of extinct strepsipterans (e.g., Grimaldi et al., 2005;Bravo et al., 2009), but the group has never been formally established under theICZN. Thus, we herein establish the family and incorporate this group into thenewly expanded key. The family includes only the species Cretostylops engeliGrimaldi and Kathirithamby, from the mid-Cretaceous deposits of northernMyanmar (Grimaldi et al., 2005).A few additional changes are incorporated herein. Protoelencholacidae waserected by Pohl & Beutel (2005) to include the fossil Protoelencholax schleeiKinzelbach (1979), but the differences given – slender mandibles with medially-crossing, blade-like apices and two tarsomeres – are typical characters of Elenchidaeand do not sufficiently distinguish the group. We therefore retain this species withinElenchidae. Also, based on the results of a recent molecular analysis, Callipharix-enidae was placed in the Halictophagidae and Lychnocolacidae was recognized as afamily distinct from the Myrmecolacidae (McMahon et al., 2011). These alterationsof the higher classification are also incorporated herein.Throughout, groups known only as fossils are denoted with a dagger ({) and otherfamilies that are also recorded from the fossil record are mentioned as such. Thecurrent classification of the order is summarized in Table 1.1 Department of Zoology, Oxford University, South Parks Road, Oxford OX1 3PS, United Kingdom(jeyaraney.kathirithamby@zoo.ox.ac.uk).2 Division of Entomology, Natural History Museum, and Department of Ecology & EvolutionaryBiology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66045 (msengel@ku.edu).3 Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79thStreet, New York, New York 10024-5192 (mengel@amnh.org).Accepted 17 June 2014; Revised 17 June 2014E 2014 Kansas Entomological SocietyJOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY87(4), 2014, pp. 385–388Key to Families of Strepsiptera(Adult males, living & fossil)1. Tarsi 5 segmented, pretarsus with pair of strong claws . . . . . . . . . . . . . . 2—. Tarsi 2–5 segmented; pretarsus with neither claws nor sensory spots; ortarsi 5-segmented and pretarsus with a pair of very weak claws, or asingle claw and with sensory spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62(1). Antenna 8 segmented; mandibles large and robust, triangular shaped,with broad base and of generalized chewing structure. . . . . . . . . . . . . . . 3—. Antenna 6–7 segmented; mandibles, if present, small, narrow, and blade-like. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53(2). Galeal lobe at base of maxillary palp present . . . . . . . . . . . {Protoxenidae—. Galeal lobe at base of maxillary palp absent . . . . . . . . . . . . . . . . . . . . . 44(3). Protrochanter and profemur free . . . . . . . . . . . {Cretostylopidae, n. fam.—. Protrochanter and profemur fused (Host: unknown). . . . . . . Bahiaxenidae5(2). Prementum free, with short palps; hind wing CuA1 short, extendingabout to middle of wing; front branch of MA absent; antenna 7-segmented, with flabella on antennomeres III and IV . . . . . . . {Mengeidae—. Prementum fused to hypopharynx; hind wing CuA1 extending almost toedge of wing; MA strongly developed with anterior branch; antenna 6-segmented, with flabella on antennomeres III to IV or III to V (Host:Zygentoma: Lepismatidae) . . . . . . . . . . . . . . . . . . . . . . . . .Mengenillidae6(1). Mandibles absent; tarsi 4–5 segmented, pretarsus with a pair of claws, ora single claw, or tarsi 4 segmented and pretarsus without claws; antenna5–7 segmented, with flabella on antennomeres III–IV or III–V (Host:Hemiptera: Cydnidae, Blissidae, Pentatomidae, Lygaeidae, Coreidae)(also known from fossils) . . . . . . . . . . . . . . . . . . . . . . . . . . Corioxenidae—. Mandibles present, narrow and blade-like; tarsi 2–4 segmented,pretarsus without claws; antenna 4–7 segmented, with flabella onantennomeres only on III or IV, or III & IV, or III to V, or III, V, & VI,or III to VI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Table 1. Hierarchical classification of Strepsiptera.Order Strepsiptera KirbyFamily {Protoxenidae Pohl et al.Family {Cretostylopidae Kathirithamby & EngelSuborder Mengenillidia KinzelbachFamily {Mengeidae PierceFamily Bahiaxenidae Bravo et al.Family Mengenillidae HofenederSuborder Stylopidia KinzelbachFamily Corioxenidae Kinzelbach (+1 fossil species)Infraorder Stylopiformia KinzelbachFamily Myrmecolacidae Saunders (+13 fossil species)Family Lychnocolacidae BohartFamily Stylopidae Kirby (+2 fossil species)Family Xenidae SaundersFamily Bohartillidae Kinzelbach (+3 fossil species)Family Elenchidae Perkins (+1 fossil species)Family Halictophagidae Perkins386 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY7(6). Tarsi 3 segmented (Host: Hemiptera: Cercopidae, Cicadellidae, Cor-eidae, Delphacidae, Eurybrachyidae, Flatidae, Fulgoridae, Issidae,Membracidae, Pentatomidae, Tettigometridae; Diptera: Tephritidae;Orthoptera: Tridactylidae; Blattaria: Blattellidae) . . . . . . . Halictophagidae—. Tarsi 2 or 4 segmented. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88(7). Tarsi 2 segmented; antenna 4 segmented (Host: Hemiptera: Delphacidae,Dictyopharidae Eurybrachidae, Flatidae, Fulgoridae, Ricaniidae). . .Elenchidae—. Tarsi 4 segmented; antenna 5–7 segmented . . . . . . . . . . . . . . . . . . . . . . 99(8). Antennae 7 segmented, with flabella on antennomeres III, V, and VI;maxillary base five times longer than palpus (Host: unknown) (alsoknown from fossils) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bohartillidae—. Antennae 4–7 segmented, with flabellum on antennomere IIII only;maxillary base equal to, or smaller than palpus . . . . . . . . . . . . . . . . . . 1010(9). Antenna 4 or 6 segmented, with flattened flabella . . . . . . . . . . . . . . . . 11—. Antenna 7 segmented, with rounded flabella . . . . . . . . . . . . . . . . . . . . 1211(10). Antenna 6 segmented, metathorax with spoon-shaped sclerite (Host:Hymenoptera: Andrenidae, Colletidae, Halictidae) (also known fromfossils) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stylopidae—. Antenna 4 segmented, metathorax with no spoon-shaped sclerite (Host:Hymenoptera: Masaridae, Mutillidae, Sphecidae, Vespidae) . . . . . Xenidae12(10). Wing venation with CuA1 and CuA2 present (Host: unknown) . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lychnocolacidae—. Wing venation with only CuA1 (Host: Males – Hymenoptera:Formicidae: Dolichoderinae, Ecitoninae, Formicinae, Myrmicinae,Pseudomyrmecinae, Ponerinae / Females – Orthoptera, Mantodea) (alsoknown from fossils) . . . . . . . . . . . . . . . . . . . . . . . . . . . . MyrmecolacidaeSystematic PaleontologyCretostylopidae Kathirithamby & Engel, new familyTYPE GENUS: Cretostylops Grimaldi and Kathirithamby in Grimaldi et al. (2005).DESCRIPTION: Male. Body size small (ca. 1.5 mm, much smaller than the massiveProtoxenos Pohl et al. which is ca. 7 mm); antenna 8-segmented, with 6 flabella onantennomeres III–VIII (a plesiomorphic condition for Strepsiptera, similar to that ofProtoxenos); bases of antennae situated between compound eyes; ommatidia small,not separated by microtrichia; mandibles robust, sclerotized, large, and triangular (adistinctive plesiomorphy shared with Protoxenos, and unlike the mandibles inBahiaxenos which are large but not as robust as other Recent Strepsiptera in whichthey are narrow, small, and blade-like); maxillary lobe lacking galeal lobe (as in allother Strepsiptera except Protoxenos); labrum reduced (also present in Protoxenosbut absent in all other strepsipterans except Mengea Grote and the AustralianMengenilla Hofeneder); tarsi long and pentamerous; pretarsus with a pair of claws,setae absent (present in Protoxenos and all basal extant groups such as Bahiaxenidaeand Mengenillidae); pseudohalters (forewings) large; hind wings with well-developedR1 and R4, with pterostigma-like thickening; protrochanter and profemur free(unlike all extant Strepsiptera).Female. Unknown.VOLUME 87, ISSUE 4 387COMMENTS: The earliest known Strepsiptera and the first described from theMesozoic is C. engeli (Grimaldi et al., 2005; Grimaldi and Engel, 2005), preserved inmid-Cretaceous amber from Myanmar. All phylogenetic analyses which includedboth fossil and modern families recover the extinct genus Protoxenos, in middleEocene Baltic amber, as sister to all other Strepsiptera, with Cretostylops the nextlineage to diverge (Grimaldi et al., 2005; Pohl and Beutel, 2005; Bravo et al., 2009;Hünefield et al., 2011). Although Cretostylops is older (99 Ma) than Protoxenos(45 Ma), the latter is clearly supported as the most basal group and must have beenan ancient relic lineage even by the Paleogene.AcknowledgementsWe are grateful for the input from two anonymous reviewers and Roy J.Beckemeyer. This is a contribution of the Division of Entomology, University ofKansas Natural History Museum.Literature CitedBravo, F., H. Pohl, A. Silva-Neto, and R. G. Beutel. 2009. Bahiaxenidae, a ‘‘living fossil’’ and a newfamily of Strepsiptera (Hexapoda) discovered in Brazil. Cladistics 25(6):614–623.Grimaldi, D., and M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press, Cambridge,UK. xv+755 pp.Grimaldi, D., J. Kathirithamby, and V. Schawaroch. 2005. Strepsiptera and triungula in Cretaceousamber. Insect Systematics and Evolution 36(1):1–20.Hünefield, F., H. Pohl, B. Wipfler, F. Beckmann, and R. G. Beutel. 2011. The male postabdomen andgenital apparatus of Mengea tertiaria, a strepsipteran amber fossil (Insecta). Journal of ZoologicalSystematics and Evolutionary Research 49(4):298–308.Kinzelbach, R. K. 1979. Das erste neotropische Fossil der Fächerflügler (Stuttgarter Bernsteinsammlung:Insecta, Strepsiptera). Stuttgarter Beiträge zur Naturkunde, Serie B (Geologie und Paläontologie)53:1–14.McMahon, D. P., A. Hayward, and J. Kathirithamby. 2011. The first molecular phylogeny of Strepsiptera(Insecta) reveals an early burst of molecular evolution correlated with the transition toendoparasitism. PLoS ONE 6:e21206 [1–10].Pohl, H., and R. G. Beutel. 2005. The phylogeny of Strepsiptera (Hexapoda). Cladistics 21(4):328–374.Pohl, H., R. G. Beutel, and R. Kinzelbach. 2005. Protoxenidae fam. nov. (Insecta, Strepsiptera) fromBaltic amber – a ‘missing link’ in strepsipteran phylogeny. Zoologica Scripta 34(1):57–69.388 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

A revised key to the living and fossil families of Strepsiptera, with the description of a new family, Cretostylopidae

Jeyaraney Kathirithamby

Michael S. Engel

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A Revised Key to the Living and Fossil Families of Strepsiptera, with the Description of a New Family, Cretostylopidae

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A revised key to the living and fossil families of Strepsiptera, with the description of a new family, Cretostylopidae

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