Hymenoptera

Larva of horned hickory sawfly Macroxyela, photographed by Charley Eiseman.

Belongs within: Holometabola.
Contains: Tenthredinoidea, Siricoidea, Vespina.

The Hymenoptera include the wasps and their derivative taxa such as bees and ants. Members of the group are primitively characterised by the presence of hamuli, small hooks on the anterior margin of the hind wing that hook onto the fore wing and hold the wings locked together. The earliest known members of the Hymenoptera in the Late Triassic are assigned to the Xyelidae, though it is uncertain whether all members of the fossil subfamily Archexyelinae form a clade with modern xyelids (Grimaldi & Engel 2005). Larvae of the Xyelidae are herbivorous, with Xyelinae feeding on developing cones or shoots of conifers, and Macroxyela on leaves of elms or walnuts. The Xyelidae have been regarded in the past as the sister group of all other Hymenoptera but a molecular phylogenetic analysis of the order by Peters et al. (2017) united them in a clade Eusymphyta with the other ectophytophagous wasp groups Pamphilioidea and Tenthredinoidea. The Pamphilioidea are a relatively small Holarctic group of sawflies with an enlarged, almost prognathous, head that is widest near the clypeus.

The remaining Hymenoptera formed a clade Unicalcarida combining the carnivorous wasps of the Vespina with the endophytophagous groups Xiphydriidae, Siricoidea and Cephoidea, whose larvae develop in burrows within their plant hosts.

Synapomorphies (from Grimaldi & Engel 2005): Hamuli present; protibial spur with velum; haplodiploid sex determination.

Hymenoptera (see below for synonymy)
    |--Eusymphyta PK17
    |    |--Pamphilioidea [Megalodontoidea] PK17
    |    |    |  i. s.: Parapamphiliidae GE05
    |    |    |         Acantholyda hieroglyphica PK17
    |    |    |--Pamphiliidae R02b
    |    |    |    |--Atocus Z02b
    |    |    |    |--Pamphilius Latreille 1802 L02
    |    |    |    |    `--*P. sylvaticus (Linnaeus 1758) L02, L58 [=Tenthredo sylvatica L02]
    |    |    |    |--Cephalcia abietis HR11
    |    |    |    `--Onycholyda amplecta HR11
    |    |    `--+--Xyelydidae R02b
    |    |       `--+--Praesiricidae R02b
    |    |          `--Megalodontes Latreille 1802 L02 [Megalodontesidae R02b]
    |    |               |--*M. cephalotes [=Tenthredo cephalotes] L02
    |    |               `--M. spireae MS01
    |    `--+--Tenthredinoidea PK17
    |       `--Xyelidae [Archihymenoptera, Xyeloidea] PK17
    |            |  i. s.: Spathoxyela pinicola R02b
    |            |         Angaridyela vitimica GE05
    |            |         Eoxyela atra R02a
    |            |         Megaxyela gigantea Mocsary 1909 Z02b, I92
    |            |         Angarixyela NR13
    |            |--Xyela [Xyelinae, Xyelini] R02a
    |            |    |--X. alpigena PK17
    |            |    `--X. julii HR11
    |            |--Macroxyela [Macroxyelinae] NPH04
    |            |    `--M. ferruginea HR11
    |            `--Archexyelinae GE05
    |                 |--Archexyela crosbyi R70
    |                 `--Madygenius GE05
    `--Unicalcarida PK17
         |  i. s.: Daohugoa Rasnitsyn & Zhang 2004 [Daohugoidae] RZ04
         |           `--*D. tobiasi Rasnitsyn & Zhang 2004 RZ04
         |--+--Xiphydriidae [Xiphydrioidea] HR11
         |  |    |--Derycyrtinae N91
         |  |    |    |--Austrocyrta australiensis N91
         |  |    |    `--Derecyrta circularis HR11
         |  |    `--Xiphydriinae N91
         |  |         |--Rhysacephala N91
         |  |         `--Xiphydria Latreille 1802 L02
         |  |              |--*X. camelus [=Sirex camelus] L02
         |  |              `--X. prolongata HR11
         |  `--Siricoidea HR11
         `--+--Cephoidea GE05
            |    |--Sepulcidae GE05
            |    `--Cephidae HR11
            |         |--Mesocephus sibiricus Z02b
            |         |--Cephus Latreille 1802 L02
            |         |    |--*C. pygmaeus [=Sirex pygmaeus] L02
            |         |    |--C. cinctus PK17
            |         |    |--C. flavicornis Lucas 1849 E12
            |         |    `--C. spinipes PK17
            |         `--Hartigia Schioedte 1838 HR11, L59
            |              `--H. trimaculata HR11
            `--Vespina HR11

Hymenoptera incertae sedis:
  Stenopolybia Z02a
  Orisema uchancoi H03
  Rielia manticida A71
  Euryxyela Hong 1984 [=Xaxexis Pagliano & Scaramozzino 1990] NPH04
  Ophrynopus schauinslandi G27
  Amalthea Rafinesque 1815 BR05
  Giraudia Foerster 1868 BR05
  Janus Stephens 1835 BR05
  Dineura virididorsata (Retzius 1783) HP-W05
  Lionotus delphinalis A-B91
  Schizonotus Ratzeburg 1852 H11
  Myzina E12
    |--M. guerinii Lucas 1849 E12
    `--M. oraniensis Lucas 1849 E12
  Tarpa levaillantii Lucas 1849 E12
  Scleroderma ruficornis Lucas 1849 E12
  ‘Promachus’ Cresson 1887 nec Loew 1848 nec Stål 1875 Z01
  Myzinum quinquecinctum (Fabricius 1775) [=Tiphia quinquecincta, Sphex quinquecinctus] BM76
  Triassoxyela foveolata PK17
  Gigantoxyela quadrifurcata NR13
  Baeosomus Thomson 1891 Z93
  Diastrophus Hartig 1840 Z94
  Patasson nitens M94
  Lophyrus Latreille 1802 L02
    `--*L. pini (Linnaeus 1758) L02, L58 [=Tenthredo pini L02]

Hymenoptera [Chalastrogastra, Neohymenoptera, Siricina, Siricomorpha, Symphyta, Tenthredines, Urocerata, Vespida, Vespidea]

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[A-B91] Athias-Binche, F. 1991. Ecology and evolution of phoresy in mites. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 27–41. SPB Academic Publishing: The Hague.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[G27] Gourlay, E. S. 1927. Notes on the New Zealand wood-wasp Ophrynopus schauinslandi Ashmead. Transactions and Proceedings of the New Zealand Institute 57: 691–693.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H11] Harvey, M. S. 2011. Notes on some Old World schizomids of the genera Ovozomus and Schizomus (Schizomida: Hubbardiidae). Records of the Western Australian Museum 26 (2): 202–208.

[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311–449.

[HP-W05] Heitland, W., & H. Pschorn-Walcher. 2005. Biology and parasitoids of the peculiar alder sawfly, Platycampus luridiventris (Fallen) (Insecta, Hymenoptera, Tenthredinidae). Senckenbergiana Biologica 85 (2): 215–231.

[HR11] Heraty, J., F. Ronquist, J. M. Carpenter, D. Hawks, S. Schulmeister, A. P. Dowling, D. Murray, J. Munro, W. C. Wheeler, N. Schiff & M. Sharkey. 2011. Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[L59] Lopes, H. de S. 1959. A revision of Australian Sarcophagidae (Diptera). Studia Ent. 2 (1–4): 33–67.

[M94] May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May pp. 365–728. CSIRO Australia.

[MS01] Mocsáry, A., & V. Szépligeti. 1901. Hymenopterák [Hymenopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 121–169. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[NPH04] Nel, A., J. F. Petrulevicius & M. Henrotay. 2004. New Early Jurassic sawflies from Luxembourg: the oldest record of Tenthredinoidea (Hymenoptera: “Symphyta”). Acta Palaeontologica Polonica 49 (2): 283–288.

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[R02a] Rasnitsyn, A. P. 2002a. Subclass Scarabaeona Laicharting, 1781. The winged insects (=Pterygota Lang, 1888). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 75–83. Kluwer Academic Publishers: Dordrecht.

[R02b] Rasnitsyn, A. P. 2002b. Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linné, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[RZ04] Rasnitsyn, A. P., & H. Zhang. 2004. A new family, Daohugoidae fam. n., of siricomorph hymenopteran (Hymenoptera=Vespida) from the Middle Jurassic of Daohugou in Inner Mongolia (China). Proceedings of the Russian Entomological Society, St. Petersburg 75 (1): 12–16.

[R70] Riek, E. F. 1970. Fossil history. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 168–186. Melbourne University Press.

[Z02a] Zherikhin, V. V. 2002a. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

[Z02b] Zherikhin, V. V. 2002b. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.

[Z93] Zimmerman, E. C. 1993. Australian Weevils (Coleoptera: Curculionoidea) vol. 3. Nanophyidae, Rhynchophoridae, Erirhinidae, Curculionidae: Amycterinae, literature consulted. CSIRO Australia.

[Z94] Zimmerman, E. C. 1994. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May. CSIRO Australia.

[Z01] Zompro, O. 2001. A review of Eurycanthinae: Eurycanthini, with a key to genera, notes on the subfamily and designation of type species. Phasmid Studies 10 (1): 19–23.

Last updated: 3 October 2017.

Cimicomorpha

Damsel bug Nabis kinbergii, photographed by Denis Crawford.

Belongs within: Panheteroptera.
Contains: Reduviidae, Tingoidea, Miridae, Anthocoridae, Cimicidae, Polyctenidae.

The Cimicomorpha is a group of terrestrial bugs characterised by a reduced number of cells in the forewing membrane, and the reduction of the spermatheca to a vermiform gland (Shcherbakov & Popov 2002). The clade includes both phytophagous (Miridae, Tingoidea) and predatory (Reduviidae, Cimicoidea) representatives. The Reduvioidea usually have a powerful, three-segmented labium that curves away from the ventral surface of the head at rest (Richards & Davies 1977).

Members of the Cimicoidea have a phallus that punctures a membrane in the female abdomen, either puncturing the vaginal walls as in the Nabidae, allowing the sperm to migrate to the ovaries through the haemolymph rather than through the oviduct, or puncturing the abdomen wall externally as in Cimicidae and Anthocoridae, through a process known as traumatic insemination (Grimaldi & Engel 2005). The Nabidae, damsel bugs, are relatively long and slender and mostly larger than other cimicoids. The Velocipedidae are a group of medium-sized oval bugs found in the Oriental region (Richards & Davies 1977).

<==Cimicomorpha [Anthocoroidea]
    |--Reduvioidea CGW91
    |    |--Pachynomidae CGW91
    |    |--Reduviidae SP02
    |    `--Elasmodema [Elasmodemidae] RD77
    `--+--+--Tingoidea SP02
       |  `--Miroidea CGW91
       |       |--Miridae SP02
       |       `--Microphysidae CGW91
       `--Cimicoidea CGW91
            |  i. s.: Plokiophilidae CGW91
            |           |--Plokiophilinae R00
            |           `--Embiophila [Embiophilinae] R00
            |                `--E. africana [=Acladina africana] R00
            |         Medocostidae CGW91
            |         Electrocoris Usinger 1942 P92
            |--+--Anthocoridae G75
            |  `--+--Cimicidae G75
            |     `--Polyctenidae G75
            `--+--Velocipedidae G75
               |    |  i. s.: Velocipeda RD77
               |    |         Scotomedes RD77
               |    `--Darniopseinae SP02
               `--Nabidae G75
                    |  i. s.: Arachnocoris GE05
                    |         Reduviolus B27
                    |           |--R. biformis Bergroth 1927 B27
                    |           |--R. capsiformis [incl. R. saundersi] B27
                    |           |--R. maoricus B27
                    |           |--R. maracandicus B27
                    |           `--R. quadripunctatus Bergroth 1927 B27
                    |         Sinanabis brevipes Zhang 1986 RJ93
                    |         Himacerus apterus HW96
                    |--Prostemmatinae CGW91
                    |    |--Phorticus CGW91
                    |    |--Prostemma CGW91
                    |    |    |--P. guttula H01
                    |    |    `--P. sanguineum H01
                    |    `--Alloeorrhynchus CGW91
                    |         |--A. flavipes WP99
                    |         |--A. flavolimbatus B27
                    |         |--A. myersi Bergroth 1927 B27
                    |         `--A. queenslandicus WEE74
                    `--Nabinae CGW91
                         |--Arbela telomi CGW91
                         |--Stenonabis CGW91
                         |--Gorpis [Gorpinae] CGW91
                         |    `--G. cribraticollis Stal 1859 K08
                         `--Nabis Latreille 1802 L02
                              |  i. s.: N. brevis H01
                              |         N. capsiformis CGW91
                              |         N. ferus H01
                              |         N. flavomarginatus H01
                              |         N. guttula [=Reduvius guttula] L02
                              |         N. limbatus H01
                              |         N. myrmicoides BM76
                              |         N. rugosus BM76
                              `--N. (Tropiconabis) M83
                                   |--N. (T.) kinbergii M83
                                   `--N. (T.) maoricus M83

Cimicomorpha incertae sedis:
  Pterocimex [Pterocimicidae] GE05
  Lygaenocoris GE05
  Mesopentacoridae [Protocoridae] GE05
    |--Mesopentacoris GE05
    |--Corienta transbaicalica Popov 1990 RJ93
    `--Pallicoris firmis Lin 1986 RJ93
  Miridoides GE05
  Scutellifer GE05
  Cymocoris GE05
  Coriopachys GE05

*Type species of generic name indicated

REFERENCES

[B27] Bergroth, E. 1927. Hemiptera Heteroptera from New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 671–684.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[G75] Gross, G. F. 1975. Plant-feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera—Part I. Handbook of the Flora and Fauna of South Australia.

[HW96] Hoebeke, E. R., & A. G. Wheeler, Jr. 1996. Meligethes viridescens (F.) (Coleoptera: Nitidulidae) in Maine, Nova Scotia, and Prince Edward Island: diagnosis, distribution, and bionomics of a Palearctic species new to North America. Proceedings of the Entomological Society of Washington 98 (2): 221–227.

[H01] Horváth, G. 1901. Hemipterák [Hemiptera]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 245–274. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[K08] Kirkaldy, G. W. 1908. A catalogue of the Hemiptera of Fiji. Proceedings of the Linnean Society of New South Wales 33: 345–391, pl. 4.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[R00] Ross, E. S. 2000. Embia: Contributions to the biosystematics of the insect order Embiidina. Occasional Papers of the California Academy of Sciences 149: 1–53, 1–36.

[SP02] Shcherbakov, D. E., & Yu. A. Popov. 2002. Superorder Cimicidea Laicharting, 1781: Order Hemiptera Linné, 1758. The bugs, cicadas, plantlice, scale insects, etc. [=Cimicida Laicharting, 1781, =Homoptera Leach, 1815+Heteroptera Latreille, 1810). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 143–157. Kluwer Academic Publishers: Dordrecht.

[WP99] Walter, D. E., & H. C. Proctor. 1999. Mites: Ecology, Evolution and Behaviour. CABI Publishing: Wallingford (UK).

[WEE74] Woodward, T. E., V. F. Eastop & J. W. Evans. 1974. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 52–57. Melbourne University Press.

Last updated: 14 May 2022.

Plesiothrips

Female and male of Plesiothrips perplexus, from CSIRO.

Belongs within: Thripidae.

Plesiothrips is a genus of thrips living on grasses, characterised by a weak ovipositor in the females (Mound & Marullo 1996). Most species of the genus are native to the Americas, with Plesiothrips perplexus widespread in tropical and subtropical regions around the world.

Characters (from Mound & Marullo 1996): Head with ocellar setae I absent. Antennal segment I with pair of well developed dorsal apical setae; females with segment III smaller than IV; males with segment III very small, IV–VI greatly enlarged. Abdominal tergites sometimes with pair of weakly developed ctenidia. Ovipositor very weak, lacking marginal serrations.

<==Plesiothrips Hood 1915 MM96
    |--*P. perplexus (Beach 1897) [=Sericothrips perplexa] MM96
    |--P. aberrans (Crawford 1941) [=Taeniothrips aberrans] MM96
    |--P. aczeli De Santis 1959 MM96
    |--P. amblycauda Hood 1925 MM96
    |--P. andropogoni Watts 1934 [incl. P. watsoni Watts 1934] MM96
    |--P. ayarsi Stannard 1957 MM96
    |--P. brunneus Hood 1936 [incl. P. fawcetti Morison 1937] MM96
    |--P. frequens (Moulton 1938) [=Thrips frequens] MM96
    |--P. longicollis Hood 1954 MM96
    |--P. maculosus Hood 1954 MM96
    |--P. o Girault 1929 G29b
    |--P. octarthrus Hood 1925 MM96
    |--P. pallidipennis Hood 1957 MM96
    |--P. panicus (Moulton 1929) [=Thrips panicus] MM96
    |--P. pulcher Girault 1929 G29a
    |--P. sakagamii Kudo 1974 MM96
    |--P. sculpticollis Hood 1950 MM96
    |--P. setiventris Hood 1950 MM96
    |--P. tricolor Johansen 1976 MM96
    |--P. typhae Hood 1940 MM96
    |--P. verticalis Hood 1940 MM96
    `--P. williamsi Hood 1940 MM96

*Type species of generic name indicated

REFERENCES

[G29a] Girault, A. A. 1929a. Description of a case of lunacy in Homo and of six-legged articulates. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 262-265).

[G29b] Girault, A. A. 1929b. New pests from Australia. VI. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 266-269).

[MM96] Mound, L. A., & R. Marullo. 1996. The thrips of Central and South America: an introduction (Insecta: Thysanoptera). Memoirs on Entomology, International 6: 1-487.

Aculeata

Holotype of Bethylonymellus cervicalis, from Penney & Jepson (2014).

Belongs within: Apocrita.
Contains: Chrysididae, Bethylidae, Euaculeata, Dryinidae.

The Aculeata include the stinging wasps, the clade of Hymenoptera in which the ovipositor has been modified into a venomous sting formed from a pair of sharpened, stiffened gonapophyses, with the egg deposited through an opening at the base of the sting. The fossil Bethylonymidae are included on the basis of their overall resemblance with aculeates, though it is uncertain whether they yet possessed a true sting or not. Bethylonymids are known as compression fossils from the latest Jurassic of central Asia (Grimaldi & Engel 2005). Living aculeates have historically been divided between the Euaculeata and Chrysidoidea but recent phylogenetic analyses indicate that the latter group is paraphyletic. 'Chrysidoid' families are parasitoids and retain antennae with the same number of segments in both sexes, in contrast to the sexually dimorphic antennae of Euaculeata (Grimaldi & Engel 2005).

Members of the families Dryinidae and Embolemidae develop attached to leafhoppers of the Auchenorrhyncha, protruding from the host's abdomen in a bulging sac (thylacium). Embolemidae have the ten-segmented antennae inserted remotely from the mouth on a frontal prominence; females are often brachypterous (Naumann 1991). The Sclerogibbidae are a widespread but not diverse group of small wasps attacking webspinners (Embioptera). The Plumariidae, another small group found in drier regions of South America and southern Africa, are of unknown natural history. Males have a relatively rich wing venation with well developed accessory veins in the apical membrane whereas females are wingless (Brothers 2011).

<==Aculeata [Bethyloidea, Chrysidoidea, Cleptiosa, Vespomorpha]
    |  i. s.: Stelopolybia H79
    |           |--S. angulata H79
    |           |--S. areata H79
    |           |--S. falvipennis H79
    |           |--S. multipicta H79
    |           |--S. myrmecophila H79
    |           `--S. testacea H79
    |         Lithoserix williamsi GE05
    |         Lancepyris opertus PK17
    |--Bethylonymidae GE05
    |    |--Bethylonymellus cervicalis R02
    |    `--Bethylonymus sibiricus RJ93
    `--+--+--Chrysididae BD17
       |  `--+--Bethylidae BD17
       |     `--+--Plumalexius Brothers 2011 [Plumalexiidae] B11
       |        |    `--*P. rasnitsyni Brothers 2011 B11
       |        `--Plumariidae BD17
       |             |--+--Plumarius Philippi 1873 B11
       |             |  |    |--P. gradifrons PK17
       |             |  |    `--P. striaticeps B11
       |             |  `--Myrmecopterina Bischoff 1914 B11
       |             `--+--Myrmecopterinella Day 1977 B11
       |                `--+--Maplurius Roig-Alsina 1994 B11
       |                   |    `--M. spatulifer B11
       |                   `--+--Pluroides Diez, Roig-Alsina & Fidalgo 2010 B11
       |                      |    `--P. porteri B11
       |                      `--+--Plumaroides Brothers 1974 B11
       |                         |    |--P. andalgalensis B11
       |                         |    |--P. brothersi B11
       |                         |    `--P. tiphlus Diez 2008 B11
       |                         `--Mapluroides Diez, Fidalgo & Roig-Alsina 2007 B11
       |                              `--M. oglobini B11
       `--+--Euaculeata BD17
          `--+--Sclerogibbidae BD17
             |    |--Probethylus schwarzi R00
             |    `--Sclerogibba N91
             |         |--S. africana B11
             |         |--S. berlandi B11
             |         `--S. magrettii B11
             `--+--Dryinidae BD17
                `--Embolemidae BD17
                     |--Embolemus N91
                     |    |--E. brothersi B11
                     |    |--E. collinsi B11
                     |    |--E. confusa B11
                     |    `--E. magna B11
                     `--Ampulicomorpha succinalis RJ93

*Type species of generic name indicated

REFERENCES

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

[B11] Brothers, D. J. 2011. A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata). ZooKeys 130: 515–542.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H79] Howse, P. E. 1979. The uniqueness of insect societies: Aspects of defense and integration. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 345–374. Academic Press: London.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[R02] Rasnitsyn, A. P. 2002. Superorder Vespidea Laicharting, 1781. Order Hymenoptera Linné, 1758 (=Vespida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 242–254. Kluwer Academic Publishers: Dordrecht.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[R00] Ross, E. S. 2000. Embia. Contributions to the biosystematics of the insect order Embiidina. Occasional Papers of the California Academy of Sciences 149: 1–53, 1–36.

Last updated: 13 March 2021.

Helicteroideae

East Indian screw tree Helicteres isora, photographed by J. M. Garg.

Belongs within: Malvaceae.

The Helicteroideae is a tropical clade within the Malvaceae supported by molecular analysis. Baum et al. (1998) defined it as the most exclusive clade including Durio zibethinus and Helicteres isora. Durio zibethinus, durian, is a native of southeastern Asia whose large, thorny-skinned fruits are variously prized or reviled for their strongly scented flesh.

Characters (from the Angiosperm Phylogeny Website): Petiole pulvinate; sepals connate, petals clawed, often with lateral constrictions; androgynophore +; stamens 10-many, usually with short tube and/or fascicles; pollen baculate or microverrucate to suprareticulate; outer and inner integuments 2 cells across; n = 9, 14, 20, 25, etc.

<==Helicteroideae BAN98
    |--+--Coelostegia BAN98
    |  |--Neesia BAN98
    |  |--Cullenia BAN98
    |  |--Kostermansia BAN98
    |  `--Durio [Durioneae] BAN98
    |       |--D. graveolens K03
    |       `--D. zibethinus BAN98
    `--+--Reevesia BAN98
       |--Triplochiton scleroxylon BAN98, V01
       |--Ungeria BAN98
       |--Neoregnellia BAN98
       `--Helicteres BAN98
            |--H. angustifolia LK14
            |--H. cana LK14
            |--H. guazumifolia MM96
            |--H. isora BAN98
            `--H. rhynchocarpa LK14

*Type species of generic name indicated

REFERENCES

[BAN98] Baum, D. A., W. S. Anderson & R. Nyffeler. 1998. A durian by any other name: taxonomy and nomenclature of the core Malvales. Harvard Papers in Botany 3 (2): 315–330.

[K03] Kulip, J. 2003. An ethnobotanical survey of medicinal and other useful plants of Muruts in Sabah, Malaysia. Telopea 10 (1): 81–98.

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[MM96] Mound, L. A., & R. Marullo. 1996. The thrips of Central and South America: an introduction (Insecta: Thysanoptera). Memoirs on Entomology, International 6: 1–487.

[V01] Vánky, K. 2001. The emended Ustilaginaceae of the modern classificatory system for smut fungi. Fungal Diversity 6: 131–147.

Last updated: 22 July 2021.

Acanthaceae

Staurogyne leptocaulis ssp. decumbens, copyright Kevin Thiele.

Belongs within: Lamiales.
Contains: Thunbergioideae, Acanthoideae.

The Acanthaceae is a group of mostly tropical herbs, shrubs and vines. They are commonly divided between three subfamilies, of which the Nelsonioideae have non-explosive fruits without a retinaculum and descending-cochlear flower aestivation (Scotland & Vollesen 2000). Members of the Nelsonioideae include Staurogyne, a pantropical genus of herbs and small shrubs found in damp habitats that may be at least seasonally amphibious.

<==Acanthaceae
    |  i. s.: Multiareolites formosus Pares Regali et al. 1984 CBH93
    |         Multimarginites vanderhammeni Germeraad et al. 1968 CBH93
    |         Enslenia Raf. 1817 KC01
    |--+--Thunbergioideae OP01
    |  `--Acanthoideae OP01
    `--Nelsonioideae SV00
         |--Anisosepalum SV00
         |--Gynocraterium SV00
         |--Ophiorrhiziphyllon SV00
         |--Saintpauliopsis SV00
         |--Elytraria [incl. Tubiflora] SV00
         |    `--E. crenata SV00
         |--Nelsonia C55
         |    |--N. campestris LK14
         |    `--N. canescens C55
         `--Staurogyne [incl. Ancistrostylis, Ebermaiera, Erythracanthus, Neozenkerina, Staurogynopsis, Zenkerina] SV00
              `--S. leptocaulis LK14
                   |--S. l. ssp. leptocaulis LK14
                   `--S. l. ssp. decumbens LK14

*Type species of generic name indicated

REFERENCES

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[CBH93] Collinson, M. E., M. C. Boulter & P. L. Holmes. 1993. Magnoliophyta (‘Angiospermae’). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 809–841. Chapman & Hall: London.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby's Dictionary of the Fungi 9^th ed. CAB International: Wallingford (UK).

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[OP01] Olmstead, R. G., C. W. dePamphilis, A. D. Wolfe, N. D. Young, W. J. Elisons & P. A. Reeves. 2001. Disintegration of the Scrophulariaceae. American Journal of Botany 88 (2): 348–361.

[SV00] Scotland, R. W., & K. Vollesen. 2000. Classification of Acanthaceae. Kew Bulletin 55 (3): 513–589.

Last updated: 3 April 2020.

Coelosphaeridae

Live individual of Forcepia, from the Florida Sea Grant.

Belongs within: Poecilosclerida.

The Coelosphaeridae is a group of sponges with reduced inhalant and exhalant openings.

Characters (from Bergquist 1978): Generally of spherical shape; inhalant and exhalant openings reduced to specialised tubular fistules; remainder of surface a dense layer of tangentially disposed diactinal megascleres. Choanosome a very delicate network of plumoreticulate spicule fibres, collapsing if sponge is removed from water. Microscleres are chelae, toxas, sigmas and microxeas.

<==Coelosphaeridae
    |--Coelosphaera Thomson 1873 [incl. Histoderma Carter 1874] BJ06
    |    |--‘Histoderma’ dichela Hentschel 1912 BJ06
    |    |    |--H. d. dichela BJ06
    |    |    `--H. d. gracilis Hentschel 1912 BJ06
    |    `--‘Histoderma’ navicelligerum BJ06
    |         |--H. n. navicelligerum BJ06
    |         `--H. n. aruensis Hentschel 1912 BJ06
    `--Forcepia Carter 1874 [incl. Wilsa de Laubenfels 1930] L01
         |  i. s.: F. mertoni Hentschel 1912 BJ06
         `--F. (Forcepia) L01
              |--F. (F.) acanthostylosa Lee 2001 L01
              |--F. (F.) elvini Lee 2001 L01
              |--F. (F.) forcipula (Lundbeck 1905) [=Esperiopsis forcipula, Leptolabis forcipula] L01
              |--F. (F.) hartmani Lee 2001 L01
              |--F. (F.) hymena (de Laubenfels 1930) [=Wilsa hymena] L01
              |--F. (F.) japonica Koltun 1959 L01
              |--F. (F.) macrostylosa Lee 2001 [=F. macrotylota (l. c.)] L01
              `--F. (F.) topsenti Lundbeck 1905 L01

*Type species of generic name indicated

REFERENCES

[BJ06] Barnich, R., & D. Janussen. 2006. Die Typen und Typoide des Naturmuseums Senckenberg, Nr. 86. Type catalogue of the Porifera in the collections of the Senckenberg Museum in Frankfurt am Main, Germany. Senckenbergiana Biologica 86 (2): 127-144.

Bergquist, P. R. 1978. Sponges. University of California Press.

[L01] Lee, W. L. 2001. Four new species of Forcepia (Porifera, Demospongiae, Poecilosclerida, Coelosphaeridae) from California, and synonymy of Wilsa de Laubenfels, 1930, with Forcepia, Carter, 1874. Proceedings of the California Academy of Sciences 52 (18): 227-244.