Vespidae

Female Parischnogaster striatula provisioning a larval cell, copyright David Baracchi.

Belongs within: Euaculeata.
Contains: Polybia, Masarinae, Eumeninae, Zethinae, Polistinae, Vespinae, Eustenogaster, Liostenogaster.

The Vespidae is a family of often large wasps including the social wasps and their close solitary or subsocial relatives. Eusocial wasps are placed in the subfamilies Vespinae, Polistinae and Stenogastrinae, with species of the last being facultatively eusocial. Stenogastrine colonies, usually composed of up to ten individuals, are flexible in structure and they lack the true workers of other social wasps (Hines et al. 2007). Members of the Stenogastrinae are characterised by modification of the first metasomal segment into an extremely long petiole, as long as the remaining metasomal segments together. They are sometimes referred to as 'hover wasps' owing the characteristic hovering behaviour engaged in by species of some genera. Solitary wasps include the distinctive genus Euparagia, restricted to southwest North America, of which the species E. scutellaris is known to provision its nests with weevils.

Characters (from Naumann 1991): Mostly large (5-32 mm). Inner margins of compound eyes strongly emarginate except in Masarinae; mandibles generally short and broad but becoming elongate and crossing to form a distinct 'X' in many Eumeninae; glossae and paraglossae sometimes greatly produced and with apical sclerotised pads (acroglossal buttons); pronotum not freely movable against mesothorax, mid-dorsally short, posterolateral corner acutely produced dorsal to tegula and spiracle cover lobe close to tegula (remote in Stenogastrinae); parategula (an apically rounded, more or less dorsoventrally flattened lobe arising from posterolateral corner of mesoscutum and lying adjacent to tegula) present in Eumeninae and some Masarinae; calcar of hind tibia apically curved, bifid or trifid in some Masarinae, strigil well developed; fore wing at rest folded longitudinally in all except Stenogastrinae, Euparagiinae and many Masarinae; primitively with three submarginal cells in fore wing; cell 1M of fore wing elongate; hind wing jugal lobe often present; terga primitively overlapping sterna and freely movable against them, but with various degrees of fusion in many taxa; posterior metasomal segments often retractile within segment 2; T3 and S3 often enlarged; females ovipositing into cells before provisioning. Larvae variable, usually spinning a cocoon.

<==Vespidae [Eumenidae, Eumenididae, Vespariae]
    |--+--+--Masarinae HH07
    |  |  `--Euparagia Cresson 1879 HH07, S86 [Euparagiinae]
    |  |       |--E. richardsi HH07
    |  |       `--E. scutellaris WB03
    |  `--+--Eumeninae HH07
    |     `--+--Zethinae HH07
    |        `--+--Polistinae HH07
    |           `--Vespinae HH07
    `--Stenogastrinae HH07
         |  i. s.: Metischnogaster van der Vecht 1977 NK11
         |           |--M. cillipennis (Smith 1857) NK11
         |           `--M. drewseni (de Saussure 1857) NK11
         |         Cochlischnogaster Dong & Otsuka 1997 NK11
         |           `--C. spatulata Carpenter & Starr 2000 NK11
         |--+--Eustenogaster HH07
         |  `--Liostenogaster HH07
         `--Parischnogaster von Schulthess 1914 HH07, NK11
              |--P. alternata Sakagami 1969 NK11
              |--P. aurifrons (Smith 1862) NK11
              |--P. foveata (du Buysson 1907) NK11
              |--P. gracilipes (van der Vecht 1977) NK11
              |--P. jacobsoni (du Buysson 1913) NK11
              |--P. mellyi (de Saussure 1852) NK11
              |--P. nigricans (Cameron 1902) NK11
              |--P. striatula (du Buysson 1907) NK11
              `--P. unicuspidata Reyes 1988 NK11

Vespidae incertae sedis:
  Sinostigma Hong 1983 E01
    `--*S. spinalata Hong 1983 E01
  Curiosivespa R02
  Agelaia multipicta BR07
  Odontodynerus GS61
    |--O. brincki Giordani Soika 1961 GS61
    |--O. quadrituberculatus (Smith 1857) [=Odynerus quadrituberculatus] GS61
    |--O. rikatlensis [incl. O. hansi] GS61
    `--O. silaos (Saussure 1853) [=Odynerus silaos; incl. Ody. silaensis Saussure 1855] GS61
  Afreumenes GS61
    |--A. melanosoma GS61
    `--A. violaceus (Giordani Soika 1941) [=Eumenes (Afreumenes) violaceus] GS61
  Leptomenes GS61
    |--L. subtilis Giordani Soika 1938 GS61
    `--L. vulneratus (Saussure 1855) (see below for synonymy) GS61
  Quartinia capensis [incl. Q. scutellimacula Schulthess 1929, Q. capensis var. scutellimacula] GS61
  ‘Didineis’ solidescens Scudder 1890 BM76
  Epipona Latreille 1802 L02
    |--E. nidulans [=Vespa nidulans] L02
    `--E. tatua (Cuvier 1797) L02, C97 [=Vespa tatua L02]
  Nectarina B06
    |--N. augusti [incl. N. augusti var. quinta] B06
    `--N. lecheguana B06
  Chartergus B06
    |--C. apicalis G74
    |--C. chartarius H79
    `--C. globiventris B06
  Parachartergus bentobuenoi B06
  Polybia B06
  Synaeca B06
    |--S. cyanea B06
    `--S. surinama B06
  Leipomeles lamellaria B06
  Megacanthopus surinamensis B06
  Priorvespa bullata Carpenter & Rasnitsyn 1990 RJ93
  Leptochilus ornatus [incl. Stenodynerus zendaloides] S96
  Hoplomerus R13
    |--H. melanocephalus R13
    |--H. reniformis R13
    `--H. spinipes R13

Leptomenes vulneratus (Saussure 1855) [=Odynerus vulneratus; incl. Labus transiens Kohl 1907, Leptomenes transiens] GS61

*Type species of generic name indicated

REFERENCES

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

[B06] Brèthes, J. 1906. Véspidos y eumenídidos sudamericanos (nuevo suplemento). Anales del Museo Nacional de Buenos Aires, serie 3, 6: 310–377.

[BR07] Buzatto, B. A., G. S. Requena, E. G. Martins & G. Machado. 2007. Effects of maternal care on the lifetime reproductive success of females in a Neotropical harvestman. Journal of Animal Ecology 76 (5): 937–945.

[C97] Cuvier, G. 1797. Note sur une nouvelle espèce de guépe cartonnière. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (8): 57–58.

[E01] Engel, M. S. 2001. A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History 259: 1–192.

[GS61] Giordani Soika, A. 1961. Hymenoptera (Aculeata): Vespidae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 440–451. Almqvist & Wiksell: Uppsala.

[G74] Günther, K. K. 1974. Staubläuse, Psocoptera. Die Tierwelt Deutschlands 61: 1–315.

[HH07] Hines, H. M., J. H. Hunt, T. K. O'Connor, J. J. Gillespie & S. A. Cameron. 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences of the USA 104 (9): 3295–3299.

[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.

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

[NK11] Nicholas, A. S., J. Kojima, M. Y. Ruslan, M. Fauzi & A. B. Idris. 2011. A list of social vespid wasps (Hymenoptera: Vespidae) from Kuala Lompat, Krau Wildlife Reserve, Pahang, peninsular Malaysia. Serangga 16 (1): 59–74.

[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.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[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.

[S96] Smith, D. R. 1996. Review of the Gasteruptiidae (Hymenoptera) of eastern North America. Proceedings of the Entomological Society of Washington 98 (3): 491–499.

[S86] Snelling, R. R. 1986. The taxonomy and nomenclature of some Australian paragiine wasps (Hymenoptera: Masaridae). Contributions in Science 378: 1–19.

[WB03] Ward, P. S., & S. G. Brady. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361–386.

Last updated: 19 May 2020.

Camponotus

Camponotus maritimus, from Antwiki.org.

Belongs within: Camponotini.
Contains: Camponotus (Camponotus), Camponotus (Myrmamblys), Camponotus (Tanaemyrmex).

Camponotus is a diverse and cosmopolitan genus of ants, generally with a characteristic rounded profile to the rear of the mesosoma. They are commonly referred to as carpenter ants owing to the habit of some species of building nests in galleries in dead wood; however, not all species in the genus exhibit this behaviour. Workers of most Camponotus species are polymorphic, with distinct major and minor forms present (some species may also possess media forms). Major workers typically have a much more massive head than minors, with larger mandibles.

Species have been divided between several subgenera but many of the resultant groupings are poorly defined and require formal phylogenetic testing; as a result, many authors eschew their usage. Phasmomyrmex, a small group of Afrotropical species with rectangular or trapezoidal heads and monomorphic workers, has historically been treated as a distinct genus but was reclassified as a subgenus of Camponotus by Ward et al. (2016). Pseudocolobopsis, the false cork-head ants, is a Neotropical group of which the majors and queens have an elongate, often anteriorly obliquely truncate head. Forelophilus is a group of small, blackish ants found in south-east Asia with a strong transverse ridge across the propodeum.

Characters (from Heterick 2009): Antenna with twelve segments (including scape); propodeal angle (if present) rounded; metapleural gland opening inconspicuous, without fringe of long setae; first gastral tergite much less than half total length of gaster; major and minor worker castes, at least, always present, media workers often present.

Camponotus Mayr 1861 T99 [incl. Myrmophyma Forel 1912 TB85, Thlipsepinotus Santschi 1928 TB85]
    |--+--C. gibbinotus Forel 1902 WBF16, TB85
    |  `--C. (Phasmomyrmex) [incl. Myrmacantha, Myrmorhachis] WBF16
    |       |--C. (P.) buchneri [incl. *Phasmomyrmex sericeus] WBF16
    |       |    |--C. b. buchneri WBF16
    |       |    `--C. b. camerounensis Ward, Blaimer & Fisher 2016 [=Phasmomyrmex buchneri griseus (preoc.)] WBF16
    |       |--C. (P.) aberrans WBF16
    |       |--C. (P.) paradoxus WBF16
    |       |    |--C. p. paradoxus WBF16
    |       |    `--C. p. cupreus WBF16
    |       `--C. (P.) wolfi WBF16
    `--+--C. (Pseudocolobopsis) WBF16
       |    |--C. (P.) claviscapus WBF16
       |    `--C. (P.) cordincola [=C. (Colobopsis) cordincola] WBF16
       `--+--+--C. hyatti WBF16
          |  `--C. maritimus WBF16
          `--+--C. bedoti WBF16
             `--C. (Forelophilus) WBF16
                  |--C. (F.) javaensis Ward, Blaimer & Fisher 2016 [=*Forelophilus overbecki (preoc.)] WBF16
                  |--C. (F.) philippinensis WBF16
                  `--C. (F.) stefanschoedli WBF16

Camponotus incertae sedis:
  C. abdominalis CM07
  C. adami Forel 1910 TB85
  C. aeneopilosus Mayr 1862 TB85
    |--C. a. aeneopilosus TB85
    `--C. a. flavidopubescens Forel 1902 TB85
  C. afflatus Viehmeyer 1925 [=C. (Myrmosaga) afflatus] TB85
  C. albivillosus Zhou 2001 Z01
  C. angusticollis (Jerdon 1851) (see below for synonymy) B03
  C. arcuatus Mayr 1876 TB85
    |--C. a. arcuatus TB85
    `--C. a. aesopus Forel 1907 TB85
  C. armstrongi McAreavey 1949 TB85 [=C. (Myrmogonia) armstrongi WBF16]
  C. auratiacus Zhou 2001 Z01
  C. auriventris Emery 1889 B03
  C. aurocinctus (Smith 1858) [=Formica aurocincta] TB85
  C. baldaccii ES12
  C. barbatus Roger 1863 B03
  C. basalis Smith 1878 [incl. C. maculatus lobinieri Forel 1902] B03
  C. bifossus [=C. (Colobopsis) bifossus] WBF16
  C. bigenus Santschi 1919 [=C. (Myrmocamelus) bigenus] TB85
  C. binghami Forel 1894 B03
  C. branneri P92
  C. breviscapus Zhou 2001 Z01
  C. buddhae Forel 1892 B03
  C. cameratus Viehmeyer 1925 [=C. (Myrmogonia) cameratus] TB85
  C. capito Mayr 1876 [=C. (*Myrmophyma) capito] TB85
    |--C. c. capito TB85
    `--C. c. ebeninithorax Forel 1915 [=C. (Myrmophyma) capito ebeninithorax] TB85
  C. carbonarius B03
  C. carin Emery 1889 [=C. dorycus carin] B03
  C. ceriseipes Clark 1938 [=C. (Myrmophyma) ceriseipes] TB85
  C. chalceoides Clark 1938 [=C. (Myrmophyma) chalceoides] TB85
  C. chalceus Crawley 1915 [=C. (Myrmosaga) chalceus] TB85
  C. cinereus Mayr 1876 TB85
    |--C. c. cinereus TB85
    |--C. c. amperei Forel 1913 [=C. (Myrmocamelus) cinereus amperei] TB85
    `--C. c. notterae Forel 1907 TB85
  C. clarior Forel 1902 H09, TB85 [=C. nigriceps clarior TB85]
  C. claripes Mayr 1876 [=C. (*Thlipsepinotus) claripes] TB85
    |--C. c. claripes TB85
    |--C. c. elegans Forel 1902 TB85
    |--C. c. inverallensis Forel 1910 TB85
    |--C. c. marcens Forel 1907 TB85
    |--C. c. minimus Crawley 1922 [=C. (Myrmophyma) claripes minima] TB85
    |--C. c. nudimalis Forel 1913 TB85
    |--C. c. orbiculatopunctatus Viehmeyer 1925 [=C. (Myrmophyma) claripes orbiculatopunctatus] TB85
    `--C. c. piperatus Wheeler 1933 [=C. (Myrmophyma) claripes piperatus] TB85
  C. compressus (Fabr. 1787) B03 (see below for synonymy)
  C. confucii Forel 1894 B03
  C. consectator (Smith 1858) [=Formica consectator] TB85
  C. consobrinus (Erichson 1842) [=Formica consobrina; incl. C. dimidiatus Roger 1863] TB85
  C. cowlei Froggatt 1896 TB85
  C. coxalis E89
  C. crassisquamis Forel 1902 B03
  C. crassus MC05
  C. crenatus Mayr 1876 TB85
  C. cruentatus (Latreille 1802) [=Formica cruentata] TB85
    |--C. c. cruentatus TB85
    `--C. c. aspera Menozzi 1925 [=C. (Myrmosericus) cruentatus aspera] TB85
  C. darlingtoni Wheeler 1934 H09, TB85 (see below for synonymy)
  C. denticulatus Kirby 1896 TB85
  C. dichrous Forel 1879 [=C. maculatus dichrous; incl. C. dichrous var. kattensis] B03
  C. discors Forel 1902 [=C. maculatus discors] TB85
    |--C. d. discors H09
    |--C. d. angustinodus (n. d.) H09
    `--C. d. yarrabahensis Forel 1915 [=C. (Myrmoturba) discors yarrabahensis] TB85
  C. dolendus Forel 1892 [=C. rufoglaucus r. dolendus] Z01
  C. donnellani H09
  C. dorycus (Smith 1860) [=Formica dorycus] TB85
    |--C. d. dorycus TB85
    `--C. d. confusus Emery 1887 TB85
  C. dromas Santschi 1919 [=C. (Myrmocamelus) dromas] TB85
  C. dryandrae H09
  C. eremicus Wheeler 1915 [=C. (Myrmogonia) eremicus] TB85
  C. erythropus Viehmeyer 1925 [=C. (Myrmosaga) erythropus] TB85
  C. esau Forel 1915 [=C. (Myrmocamelus) esau] TB85
  C. eugeniae A61
  C. evae Forel 1910 TB85
    |--C. e. evae TB85
    `--C. e. zeuxis Forel 1915 [=C. (Myrmogonia) evae zeuxis] TB85
  C. exasperatus E89
  C. extensus Mayr 1876 TB85
  C. festinus (Smith 1857) [=Formica festina] B03
  C. fictor Forel 1902 [=C. (Colobopsis) fictor] TB85
    |--C. f. fictor TB85
    `--C. f. augustulus Viehmeyer 1925 [=C. (Colobopsis) fictor augustulus] TB85
  C. fieldeae Forel 1902 TB85
  C. fieldellus Forel 1910 TB85
  C. floridanus PK17
  C. fornaronis Forel 1892 [=C. mistura fornaronis] B03
  C. froggatti Forel 1902 TB85
  C. fuscivillosus Z01
  C. gouldianus Forel 1922 TB85
  C. gretae Forel 1902 B03
  C. haberei T99
  C. hartogi Forel 1902 [incl. C. (Myrmosaga) ferruginipes Crawley in Poulton & Crawley 1922] TB85
  C. herculeanus MS01
  C. holosericeus Emery 1889 B03
  C. horni Kirby 1896 TB85
  C. inflatus Lubbock 1880 [incl. C. (Myrmamblys) aurofasciatus Wheeler 1915] TB85
  C. infuscus Forel 1892 [=C. maculatus infuscus] B03
  C. innexus Forel 1902 TB85
  C. insipidus Forel 1893 (n. d.) TB85
  C. integer [=C. sericeus var. integer] B03
  C. intrepidus (Kirby 1818) [=Formica intrepida] TB85
    |--C. i. intrepidus [incl. Formica agilis Smith 1858, C. magnus Mayr 1862] TB85
    `--C. i. bellicosus Forel 1902 TB85
  C. invidus Forel 1892 B03
  C. irritans (Smith 1857) [=Formica irritans, C. maculatus irritans; incl. C. agnatus Roger 1863] B03
  C. janeti Forel 1895 TB85
  C. janforrestae WBF16
  C. jejuensis T99
  C. jianghuaensis Xiao & Wang 1989 Z01
  C. johnclarki [=Notostigma johnclarki] H09
  C. laevigatus WB03
  C. lamarcki Forel 1892 B03
  C. largiceps Z01
  C. lasiselene Wang & Wu 1994 Z01
  C. latrunculus Wheeler 1915 [=C. (Myrmoturba) latrunculus] TB85
    |--C. l. latrunculus TB85
    `--C. l. victoriensis Santschi 1928 [=C. (Myrmoturba) latrunculus victoriensis] TB85
  C. leae Wheeler 1915 [=C. (Myrmosphincta) leae] TB85
  C. lespesi MC05
  C. lividicoxis Viehmeyer 1925 [=C. (Myrmophyma) lividicoxis] TB85
  C. longideclivis H09
  C. longifacies H09
  C. lownei Forel 1895 [=Formica nitida Lowne 1865 non Smith 1858] TB85
  C. luteus (Smith 1858) [=Formica lutea] B03
  C. macareaveyi [=C. (Myrmogonia) macareaveyi] WBF16
  C. mackayensis Forel 1902 FT08, TB85 [=C. reticulatus mackayensis TB85]
  C. marginatus (Latr. 1798) B03 (see below for synonymy)
  C. melichloros Kirby 1888 FT08
  C. melleus (Say 1835) SW07
  C. mendax Forel 1895 [=C. sericeus var. mendax] B03
  C. micans E89
  C. michaelseni Forel 1907 TB85 [incl. C. walkeri bardus Forel 1910 H09, TB85]
  C. midas Froggatt 1896 TB85
  C. minus Wang & Wu 1994 Z01
  C. mistura (Smith 1857) [=Formica mistura] B03
  C. mitis (Smith 1858) B03 (see below for synonymy)
  C. molossus Forel 1907 TB85
  C. myoporus Clark 1938 [=C. (Tanaemyrmex) myoporus] TB85
  C. nicobarensis Mayr 1865 (see below for synonymy) B03
  C. nigriceps (Smith 1858) [=Formica nigriceps] TB85
    |--C. n. nigriceps TB85
    |--C. n. lividipes Emery 1887 TB85
    |--C. n. obniger Forel 1902 TB85
    `--C. n. pallidiceps Emery 1887 TB85
  C. nigroaeneus (Smith 1858) [=Formica nigroaenea] TB85
    |--C. n. nigroaeneus TB85
    `--C. n. divus Forel 1907 TB85
  C. nirvanae Forel 1893 B03
  C. nitidiceps Viehmeyer 1925 [=C. (Myrmophyma) nitidiceps] TB85
  C. oblongus (Smith 1858) [=Formica oblonga] B03
  C. oetkeri Forel 1910 TB85
    |--C. o. oetkeri TB85
    `--C. o. voltai Forel 1913 TB85 [=C. (Myrmogonia) oetkeri voltai TB85; incl. C. rudis H09]
  C. oxleyi Forel 1902 TB85
  C. pallens E89
  C. pallidus (Smith 1857) [=Formica pallida; incl. C. rubripes subnudus Emery 1889] B03
  C. papua K91
  C. paria Emery 1889 [=C. micans paria, C. rufoglaucus paria] B03
  C. pawseyi H09
  C. pellax Santschi 1919 [=C. (Myrmocamelus) pellax] TB85
  C. perjurus H09
  C. pilicornis ES12
  C. pitjantjatarae H09
  C. postcornutus Clark 1930 [=C. (Tanaemyrmex) postcornutus] TB85
  C. prosseri H09
  C. prostans H09
  C. pseudoirritans Wu & Wang 1989 Z01
  C. punctiventris Emery 1920 [=C. (Myrmogonia) punctiventris] TB85
  C. radiatus Forel 1892 B03
  C. renggeri B14
  C. reticulatus Roger 1863 FT08 [incl. C. reticulatus var. latitans B03]
  C. rubiginosus Mayr 1876 TB85
  C. rubripes G89
    |--C. r. rubripes G89
    `--C. r. cognatus G89
  C. rufoglaucus (Jerdon 1851) [=Formica rufoglauca; incl. C. redtenbacheri Mayr 1862] B03
  C. rufus Crawley 1925 [=C. (Dinomyrmex) rufus] TB85
  C. rugipes WL04
  C. ‘sanguinea’ McAreavey 1949 (see below for synonymy) TB85
  C. scotti H09
  C. scratius Forel 1907 TB85
    |--C. s. scratius TB85
    `--C. s. nuntius Forel 1907 TB85
  C. semicarinatus (Forel 1895) [=Colobopsis rufifrons semicarinatus] TB85
  C. sericeiventris B10
  C. sericeus (Fabr. 1798) (see below for synonymy) B03
  C. siemsseni Forel 1901 B03
  C. simpsoni McArthur 2003 H09
  C. simulator Forel 1915 [=C. (Dinomyrmex) simulator] TB85
  C. spanis Xiao & Wang 1989 Z01
  C. spenceri Clark 1930 [=C. (Tanaemyrmex) spenceri, C. reticulatus Kirby 1896 non Roger 1863] TB85
  C. spinitarsus Emery 1920 [=C. (Dinomyrmex) spinitarsus] TB85
  C. sponsorum Forel 1910 TB85
  C. subnitidus Mayr 1876 TB85
    |--C. s. subnitidus TB85
    |--C. s. famelicus Emery 1887 TB85
    `--C. s. longinodis Forel 1915 [=C. (Dinomyrmex) subnitidus longinodis] TB85
  C. suffusus (Smith 1858) [=Formica suffusa] TB85
    |--C. s. suffusus [incl. Formica piliventris Smith 1858, C. schencki Mayr 1862] TB85
    `--C. s. bendigensis Forel 1902 TB85
  C. sylvaticus ES12
  C. tasmani Forel 1902 TB85
  C. taylori Forel 1892 [=C. maculatus taylori; incl. C. taylori var. infuscoides] B03
  C. terebrans (Lowne 1865) H09, TB85 [=Formica terebrans TB85]
  C. testaceipes (Smith 1858) [=Formica testaceipes] TB85
  C. thraso Forel 1893 [=C. maculatus thraso] B03
  C. tinctus (Smith 1858) E14, B03 [=Formica tincta B03]
  C. tricoloratus Clark 1941 [=C. (Tanaemyrmex) tricoloratus] TB85
  C. tristis Clark 1930 [=C. (Myrmophyma) tristis] TB85
  C. tritschleri [=C. (Colobopsis) tritschleri] WBF16
  C. tumidus Crawley 1922 [=C. (Myrmogonia) tumidus] TB85
  C. universitatis ES12
  C. varians Roger 1863 B03
  C. variegatus (Smith 1858) (see below for synonymy) Z01
  C. versicolor Clark 1930 [=C. (Myrmosaulus) versicolor] TB85
  C. villosus Crawley 1915 [=C. (Myrmoturba) villosa] TB85
  C. walkeri Forel 1893 TB85
  C. wasmanni Emery 1893 Z01 (see below for synonymy)
  C. whitei Wheeler 1915 [=C. (Myrmosphincta) whitei; incl. C. (Myrmosaulus) scutellus Clark 1930] TB85
  C. wiederkehri Forel 1894 TB85
    |--C. w. wiederkehri TB85
    `--C. w. lucidior Forel 1910 TB85
  C. wroughtoni Forel 1893 B03
  C. yerburyi Forel 1893 [=C. reticulatus yerburyi] B03
  C. (Camponotus) T99
  C. (Myrmamblys) YT99
  C. (Myrmaphaenus) ulcerosus [incl. C. bruesi] WW90
  C. (Myrmentoma) YT99
    |--C. (M.) caryae YT99
    |    |--C. c. caryae T99
    |    `--C. c. brunni T99
    |--C. (M.) decipiens WBF16
    |--C. (M.) keihitoi T99
    |--C. (M.) nipponensis T99
    |--C. (M.) quadrinotatus T99
    `--C. (M.) sayi [incl. C. rasilis] WW90
  C. (Myrmocamelus Forel 1914) E14, TB85
    |--C. (*M.) ephippium (Smith 1858) [=Formica ephippium] TB85
    |    |--C. e. ephippium TB85
    |    `--C. e. narses Forel 1915 TB85
    |--C. (M.) gambeyi Emery 1883 E14
    |    |--C. g. gambeyi E14
    |    `--C. g. marthae For. 1894 E14
    `--C. (M.) hoplites Emery 1914 E14
  C. (Myrmosaulus) camelinus (Smith 1857) N-ZLI11, B03 (see below for synonymy)
  C. (Myrmotarsus) N-ZLI11
    |--C. (M.) irritabilis N-ZLI11
    `--C. (M.) rupifemur N-ZLI11
  C. (Myrmoturba) E14
    |--C. (M.) croceomaculatus Emery 1914 E14
    |--C. (M.) maculatus (Fabricius 1781) E14, TB85 [=Formica maculata TB85, C. rubripes maculatus G89]
    |    |--C. m. maculatus TB85
    |    `--C. m. humilior Forel 1902 TB85
    `--C. (M.) novaehollandiae Mayr 1870 E14, FT08 [=C. maculatus novaehollandiae E14]
  C. (Paramyrmamblys) T99
    |--C. (P.) amamianus T99
    `--C. (P.) kiusiuensis T99
  C. (Tanaemyrmex) YT99

Camponotus angusticollis (Jerdon 1851) [=Formica angusticollis; incl. F. ardens Smith 1858, F. callida Smith 1858, F. impetuosa Smith 1858, Camponotus prismaticus Mayr 1862, C. angusticollis var. sanguinolentus] B03

Camponotus compressus (Fabr. 1787) B03 [=Formica compressa B03, C. maculatus compressus B03, C. rubripes compressus E89]

Camponotus darlingtoni Wheeler 1934 H09, TB85 [=C. (Myrmophyma) darlingtoni TB85, C. (M.) rottnesti Donisthorpe 1941 TB85]

Camponotus marginatus (Latr. 1798) B03 [=Formica marginata B03; incl. F. glabra L02, Camponotus marginatus var. himalayanus Forel 1893 B03]

Camponotus mitis (Smith 1858) B03 [=Formica mitis B03, C. maculatus mitis B03, C. rubripes mitis E89, C. (Myrmoturba) mitis Z01; incl. Formica bacchus Smith 1858 B03, C. mitis var. bacchus B03, C. mitis var. comottoi Emery 1887 B03, C. mitis var. crassinodis Forel 1892 B03, C. mitis var. dulcis Emery 1889 B03, C. variegatus var. dulcis T99, C. mitis var. fuscithorax Forel 1892 B03, F. ventralis Smith 1858 B03]

Camponotus nicobarensis Mayr 1865 [incl. C. nicobarensis exiguoguttatus Forel 1892, C. nicobarensis var. monticola Emery 1894] B03

Camponotus ‘sanguinea’ McAreavey 1949 non C. japonicus var. sanguinea Karavajew 1929 [=C. (Myrmogonia) sanguinea] TB85

Camponotus sericeus (Fabr. 1798) [=Formica sericea; incl. F. cinerascens Jerd. 1851, F. obtusa Smith 1858, Camponotus opaciventris Mayr 1878] B03

Camponotus variegatus (Smith 1858) [=Formica variegata; incl. C. (Myrmoturba) variegatus hawaiensis Emery 1920] Z01

Camponotus wasmanni Emery 1893 Z01 [=C. (Myrmepomis) wasmanni Z01, C. (Orthonotomyrmex) wasmanni Z01; incl. C. wasmanni var. mutilloides Emery 1893 B03]

Camponotus (Myrmosaulus) camelinus (Smith 1857) N-ZLI11, B03 [=Formica camelina B03; incl. F. singularis Smith 1858 B03, Camponotus (Myrmosaulus) singularis N-ZLI11]

*Type species of generic name indicated

REFERENCES

[A61] Andreae, H. 1961. Coleoptera: Cossyphodidae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 198–216. Almqvist & Wiksell: Uppsala.

[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera vol. 2. Ants and Cuckoo-Wasps. Taylor and Francis: London.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[B10] Brown, B. V. 2010. Phoridae (hump-backed flies, scuttle flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 725–761. NRC Research Press: Ottawa.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[E89] Emery, C. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XX.—Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885–87). Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 485–520.

[E14] Emery, C. 1914. Les fourmis de la Nouvelle-Calédonie & des îles Loyalty. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 4 pp. 393–436, pl. 12. C. W. Kreidels Verlag: Wiesbaden.

[ES12] Espadaler, X., & S. Santamaria. 2012. Ecto- and endoparasitic fungi on ants from the Holarctic region. Psyche 2012: 168478.

[FT08] Framenau, V. W., & M. L. Thomas. 2008. Ants (Hymenoptera: Formicidae) of Christmas Island (Indian Ocean): identification and distribution. Records of the Western Australian Museum 25 (1): 45–85.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[H09] Heterick, B. E. 2009. A guide to the ants of south-western Australia. Records of the Western Australian Museum Supplement 76: 1–206.

[K91] Kathirithamby, J. 1991. Strepsiptera. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 684–695. Melbourne University Press: Carlton (Victoria).

[L02] Latreille, P. A. 1802. Histoire Naturelle des Fourmis, et recueil de mémoires et d'observations sur les abeilles, les araignées, les faucheurs, et autres insectes. Théophile Barrois père: Paris.

[MC05] Machado, G., P. C. Carrera, A. M. Pomini & A. J. Marsaioli. 2005. Chemical defense in harvestmen (Arachnida, Opiliones): do benzoquinone secretions deter invertebrate and vertebrate predators? Journal of Chemical Ecology 31 (11): 2519–2539.

[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.

[N-ZLI11] Nur-Zati A. M., G. T. Lim & A. B. Idris. 2011. Checklist of ants of selected hill dipterocarp forests of peninsular Malaysia. Serangga 16 (1): 91–105.

[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.

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

[SW07] Schöller, M., & V. Witte. 2007. A review of the genus Clytrasoma Jacoby 1908, with description of a new species collected within a Camponotus sp. ant-nest (Insecta: Coleoptera, Chrysomelidae; Hymenoptera, Formicidae). Senckenbergiana Biologica 87 (1): 51–61.

[TB85] Taylor, R. W., & D. R. Brown. 1985. Formicoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 1–149. Australian Government Publishing Service: Canberra.

[T99] Terayama, M. 1999. The ant genus Camponotus Mayr (Hymenoptera: Formicidae) in Japan. Memoirs of the Myrmecological Society of Japan 1: 25–48.

[WL04] Wang, X.-H., & P. J. Lester. 2004. A preliminary study of the usefulness of morphometric tools for splitting the Monomorium antarcticum (Smith) complex (Hymenoptera: Formicidae), New Zealand’s most common native ants. New Zealand Entomologist 27: 103–108.

[WBF16] Ward, P. S., B. B. Blaimer & B. L. Fisher. 2016. A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinomyrmex. Zootaxa 4072 (3): 343–357.

[WB03] Ward, P. S., & S. G. Brady. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361–386.

[WW90] Wheeler, G. C., & J. Wheeler. 1990. Insecta: Hymenoptera Formicidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1277–1294. John Wiley & Sones: New York.

[YT99] Yamane, S., & M. Terayama. 1999. A new species of the genus Pristomyrmex Mayr from Japan, and a proposal of a new synonym of species in the genus Camponotus Mayr (Hymenoptera: Formicidae). Memoirs of the Myrmecological Society of Japan 1: 17–24.

[Z01] Zhou, S. 2001. Ants of Guangxi. Guangxi Normal University Press: Guilin (China).

Last updated: 25 April 2022.

Polistinae

Polybioides raphigastra, copyright Centre for Biodiversity Genomics.

Belongs within: Vespidae.
Contains: Polistes, Ropalidia.

The Polistinae, paper wasps, are a group of eusocial wasps that get their vernacular name from the usual material of their nests though they are not the only wasps to make paper nests. Reproductive queens of Polistinae are usually little distinct morphologically from the non-reproductive workers. With about 800 known species, the Polistinae are much more diverse than their sister group, the Vespinae (Hines et al. 2007). Polistines differ from vespines in not having first metasomal tergite anteriorly truncate, so the anterior and posterior sections are not strongly angled to each other, and in having an anal lobe on the hind wing (Richards 1978).

<==Polistinae
    |--+--+--Apoica pallida HH07
    |  |  `--+--Parachartergus fraternus HH07
    |  |     `--Metapolybia cingulata HH07
    |  `--+--Polistes HH07
    |     `--Mischocyttarus HH07
    |          |--M. drewseni H79
    |          |--M. flavitarsis BS06
    |          |--M. labiatus B06
    |          `--M. mexicanus HH07
    `--Ropalidiini [Ropalidiinae] HH07
         |--Ropalidia HH07
         |--Parapolybia de Saussure 1854 NK11
         |    |--P. indica (de Saussure 1854) NK11 [=Polybia indica R78]
         |    |--P. nodosa NK11
         |    `--P. varia (Fabricius 1787) NK11 [incl. Polybia orientalis R78]
         |--Belonogaster HH07
         |    |--B. brachystomus Kohl 1894 GS61
         |    |--B. filiventris (Saussure 1853) [=Raphigaster filiventris; incl. B. braunsi Kohl 1894] GS61
         |    |--B. juncea (Fabricius 1781) [=Vespa juncea; incl. B. colonialis Kohl 1894] GS61
         |    `--B. lateritius Gerstaecker 1857 GS61
         `--Polybioides du Buysson 1913 NK11
              |--P. angustus NK11
              |--P. gracilis van der Vecht 1966 NK11
              |--P. psecas du Buysson 1913 NK11
              `--P. raphigastra (de Saussure 1854) NK11 [incl. Polybia sumatrensis R78]

*Type species of generic name indicated

REFERENCES

[BS06] Brady, S. G., T. R. Schultz, B. L. Fisher & P. S. Ward. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the USA 103 (48): 18172–18177.

[B06] Brèthes, J. 1906. Véspidos y eumenídidos sudamericanos (nuevo suplemento). Anales del Museo Nacional de Buenos Aires, serie 3, 6: 310–377.

[GS61] Giordani Soika, A. 1961. Hymenoptera (Aculeata): Vespidae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 440–451. Almqvist & Wiksell: Uppsala.

[HH07] Hines, H. M., J. H. Hunt, T. K. O'Connor, J. J. Gillespie & S. A. Cameron. 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences of the USA 104 (9): 3295–3299.

[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.

[NK11] Nicholas, A. S., J. Kojima, M. Y. Ruslan, M. Fauzi & A. B. Idris. 2011. A list of social vespid wasps (Hymenoptera: Vespidae) from Kuala Lompat, Krau Wildlife Reserve, Pahang, peninsular Malaysia. Serangga 16 (1): 59–74.

[R78] Richards, O. W. 1978. The Australian social wasps (Hymenoptera: Vespidae). Australian Journal of Zoology, Supplementary Series 61: 1–132.

Last updated: 12 May 2020.

Eumeninae

Pachodynerus nasidens, copyright rjspeer.

Belongs within: Vespidae.
Contains: Delta, Paralastor, Acarodynerus, Antamenes, Stenodyneriellus, Pseudalastor, Australodynerus, Rhynchium, Discoelius, Eumenes, Pseudepipona, Symmorphus, Odynerus, Euodynerus, Montezumia, Ancistrocerus, Stenodynerus.

The Eumeninae, potter and mason wasps, are a group of vespid wasps including both solitary and presocial species (Hines et al. 2007). Members of the group are distinguished from other vespids by the presence of parategulae and bifid claws. The name 'potter wasp' refers to the pot-shaped nests built by species of Eumenes and related genera; members of this clade (the first branching in the tree below) also have the first segment of the metasoma modified into an elongate petiole (Carpenter & Cumming 1985). The remaining mason wasps, in which the first metasomal segment is generally shorter and/or broader, generally construct nests in cavities such as burrows in the ground or narrow hollows in wood. Other features uniting many members of this latter group include five-toothed mandibles, cephalic foveae in the females, and a basomedian sulcus on the second metasomal sternite (Carpenter & Cumming 1985).

<==Eumeninae [Discoeliinae, Raphiglossinae]
    |--+--Alphamenes campanulatus HH07
    |  |--Eumenes HH07
    |  |--Minixi mexicanum CC85
    |  `--Omicron CC85
    |       |--O. argentina [=Eumenes (Omicron) argentina] B06
    |       |--O. bonariensis (Brèthes 1905) [=Eumenes (Omicron) bonariensis] B05
    |       |--O. globicollis [=Eumenes (Omicron) globicollis] B05
    |       `--O. spegazzinii (Brèthes 1905) [=Eumenes (Omicrom) spegazzinii] B05
    `--+--Pseudepipona CC85
       |--+--Symmorphus HH07
       |  `--+--Pterocheilus CC85
       |     |--Odynerus CC85
       |     |--Tropidodynerus CC85
       |     |--Gymnomerus CC85
       |     `--Paravespa CC85
       |          |  i. s.: P. spinigera GS61a
       |          `--P. (Gestrodynerus Giordani Soika 1961) GS61a
       |               |--P. (*G.) gestroi [=Odynerus gestroi] GS61a
       |               |--P. (G.) minutepunctata Giordani Soika 1961 GS61a
       |               `--P. (G.) violaceipennis Giordani Soika 1961 GS61a
       `--+--+--+--Euodynerus HH07
          |  |  `--Pachodynerus Brèthes 1906 HH07, B06
          |  |       |--P. argentinus (see below for synonymy) B06
          |  |       |--P. brevithorax [=Odynerus brevithorax] B06
          |  |       |--P. erynnis HH07
          |  |       |--P. nasidens [=Odynerus nasidens] B06
          |  |       |--P. nigriculus [=Odynerus nigriculus] B06
          |  |       |--P. praecox [=Odynerus praecox] B06
          |  |       |--P. simplicicornis [=Odynerus simplicicornis] B06
          |  |       `--P. validus Brèthes 1906 B06
          |  `--+--Pseudodynerus quadrisectus HH07
          |     `--+--Paranortonia CC85
          |        `--+--Montezumia CC85
          |           `--Monobia CC85
          |                |--M. angulosa B06
          |                |    |--M. a. angulosa B06
          |                |    `--M. a. cingulata B06
          |                |--M. anisitsii Brèthes 1906 B06
          |                |--M. apicalipennis B06
          |                |--M. caridei Brèthes 1906 B06
          |                |--M. maxillaris B06
          |                `--M. quadridens OK99
          `--+--Ancistrocerus HH07
             `--+--Gastrodynerus CC85
                |--+--Smeringodynerus CC85
                |  `--Cephalodynerus CC85
                |       |--C. deformiceps CC85
                |       |--C. platycerus CC85
                |       `--C. russipes CC85
                |--+--Dolichodynerus CC85
                |  |    |--D. turgiceps CC85
                |  |    `--D. vandykei CC85
                |  `--+--Leptochiloides CC85
                |     `--+--Hypalastoroides mexicanus CC85
                |        `--+--Maricopodynerus maricoporum CC85
                |           `--Microdynerus lissomomus CC85
                `--+--Leucodynerus CC85
                   `--+--Stenodynerus HH07
                      |--Paraleptomenes miniatus CC85
                      `--Pseudonortonia CC85
                           |--*P. difformis [=Odynerus difformis] GS61b
                           |--P. androcles GS61b
                           |--P. bisuturalis [=Nortonia bisuturalis] OK99
                           `--P. luzonensis GS61b

Eumeninae incertae sedis:
  Delta C85
  Paralastor C85
  Abispa Mitchell 1838 (Abispa) [incl. Monerebia Saussure 1852] C85
    |--*A. (A.) australiana (Mitchell 1838) [=Vespa (*A.) australiana] C85
    |--A. (A.) ephippium (Fabricius 1775) [=Vespa ephippium; incl. A. meadewaldoensis Perkins 1914] C85
    |--A. (A.) laticincta van der Vecht 1960 C85
    `--A. (A.) splendida (Guérin-Méneville 1838) [=Odynerus splendidus, *Monerebia splendida] C85
         |--A. s. splendida C85
         `--A. s. australis Smith 1857 C85
  Subancistrocerus Saussure 1855 C85
    |--*S. sichelii (Saussure 1855) [=Odynerus sichelii] C85
    |--S. clavicornis GS61b
    |--S. kankauensis GS61b
    |--S. monstricornis (Giordani Soika 1941) [=Ancistrocerus (Subancistrocerus) monstricornis] C85
    |--S. palauensis [=Pseudonortonia palauensis] GS61b
    |--S. thalassorctos GS61b
    `--S. yapensis [=Pseudonortonia yapensis] GS61b
  Acarodynerus C85
  Parifodynerus Giordani Soika 1962 C85
    |--*P. parificus Giordani Soika 1962 C85
    `--P. alariformis (Saussure 1856) (see below for synonymy) C85
  Leptomenoides Giordani Soika 1962 C85
    |--*L. placidior Giordani Soika 1962 C85
    |--L. cairnensis Giordani Soika 1962 C85
    |--L. extraneus (Saussure 1855) (see below for synonymy) C85
    |--L. histrio Giordani Soika 1962 C85
    |--L. mackayensis Giordani Soika 1962 C85
    |--L. pachymeniformis Giordani Soika 1962 C85
    `--L. pronotalis Giordani Soika 1962 C85
  Antamenes C85
  Diemodynerus Giordani Soika 1962 C85
    |--*D. diemensis (Saussure 1853) [=Odynerus (Leionotus) diemensis] GS61b
    |--D. decipiens (Saussure 1867) [=Odynerus (Leionotus) decipiens, O. (Rhynchium) decipiens] C85
    |    |--D. d. decipiens C85
    |    `--D. d. positus Giordani Soika 1977 C85
    |--D. pseudacarodynerus Giordani Soika 1962 C85
    `--D. saucius (Saussure 1856) [=Odynerus saucius] C85
  Stenodyneriellus C85
  Flammodynerus Giordani Soika 1962 C85
    |--*F. subalaris (Saussure 1855) [=Odynerus subalaris] GS61b
    |--F. flammiger (Saussure 1856) [=Odynerus flammiger] C85
    |    |--F. f. flammiger C85
    |    `--F. f. nigroflammeus Giordani Soika 1962 C85
    `--F. pseudoloris Giordani Soika 1962 C85
  Pseudalastor C85
  Australodynerus C85
  Acarozumia Bequaert 1921 C85
    `--*A. amaliae (Saussure 1869) (see below for synonymy) C85
  Alastoroides Saussure 1856 (nom. cons.) [=Paralastoroides Saussure 1856 (nom. inv.)] C85
    `--*A. clotho (Lepeletier 1841) [=Alastor clotho, *Paralastoroides clotho] C85
  Allorhynchium van der Vecht 1963 C85
    |--*A. argentatum (Fabricius 1804) [=Vespa argentata] C85
    `--A. iridipenne (Smith 1861) [=Rhynchium iridipenne] C85
  Bidentodynerus Giordani Soika 1977 C85
    `--*B. bicolor (Saussure 1855) [=Odynerus bicolor, O. (Rhynchium) bicolor, Pseudepipona bicolor] C85
         |--B. b. bicolor C85
         |--B. b. aurantiopicta (Giordani Soika 1941) C85 (see below for synonymy)
         |--B. b. flavescentulus (Giordani Soika 1941) C85 (see below for synonymy)
         `--B. b. nigrocinctoides (Giordani Soika 1941) C85 (see below for synonymy)
  Ectopioglossa Perkins 1912 C85
    `--E. polita (Smith 1861) C85
         |--E. p. polita C85
         `--E. p. australensis (Meade-Waldo 1910) (see below for synonymy) C85
  Epiodynerus Giordani Soika 1958 C85
    |--*E. alecto (Lepeletier 1841) [=Odynerus alecto, Pseudepipona (*Epiodynerus) alecto] C85
    |--E. decoratus (Saussure 1855) [=Rhynchium decoratum, Pseudepipona (Epiodynerus) decorata] C85
    |--E. nigrocinctus (Saussure 1853) (see below for synonymy) C85
    |--E. tamarinus (Saussure 1853) (see below for synonymy) C85
    |    |--E. t. tamarinus C85
    |    `--E. t. inviolatus Giordani Soika 1977 C85
    `--E. tasmaniensis (Saussure 1853) C85 (see below for synonymy)
  Eudiscoelius Friese 1904 [incl. Euchalcomenes Turner 1908] C85
    |--*E. metallicus Friese 1904 C85
    `--E. gilberti (Turner 1908) [=*Euchalcomenes gilberti, Nortonia gilberti, Pachymenes gilberti] C85
  Katamenes Meade-Waldo 1910 C85
    `--K. fluctuans (Saussure 1852) [=Eumenes fluctuans] C85
  Macrocalymma Perkins 1908 C85
    |--*M. smithianum Perkins 1908 C85
    `--M. aliciae Meade-Waldo 1914 C85
  Orancistrocerus van der Vecht 1963 C85
    `--O. drewseni (Saussure 1857) [=Odynerus drewseni] C85
  Pachycoelius Giordani Soika 1969 C85
    |--*P. brevicornis Giordani Soika 1969 C85
    |--P. carinatus (Meade-Waldo 1910) [=Discoelius carinatus] C85
    `--P. mediocris Giordani Soika 1969 C85
  Ancistroceroides Saussure 1855 C85
  Parodynerus Saussure 1855 C85
    `--P. bicincta (Fabricius 1781) (see below for synonymy) C85
  Pseudabispa van der Vecht 1960 C85
    |--*P. abispoides (Perkins 1912) (see below for synonymy) C85
    |--P. confusa van der Vecht 1960 C85
    |--P. ephippioides van der Vecht 1960 C85
    `--P. paragioides (Meade-Waldo 1910) [=Abispa paragioides] C85
  Pseudozethus Perkins 1914 C85
    |--*P. australensis Perkins 1914 C85
    |--P. confusus Giordani Soika 1969 C85
    |--P. elongatus GS61b
    |--P. ephippium (Saussure 1855) [=Discoelius ephippium] C85
    |--P. insignis (Saussure 1856) [=Discoelius insignis, Macrocalymma insignis] C85
    |--P. pseudospinosus Giordani Soika 1969 [=Deuterodiscoelius pseudospinosus] C85
    |--P. spinosus (Saussure 1855) [=Discoelius spinosus] C85
    `--P. verreauxii (Saussure 1852) [=Discoelius verreauxii, Deuterodiscoelius verreauxii] C85
  Rhynchium C85
  Syneuodynerus Blüthgen 1951 C85
    |--*S. egregius (Herrich-Schaeffer 1839) [=Odynerus egregius, Euodynerus (*Syneuodynerus) egregius] C85
    |--S. aurantiopilosellus (Giordani Soika 1962) C85 (see below for synonymy)
    |--S. longebispinosus (Giordani Soika 1962) [=Pseudepipona (Syneuodynerus) longebispinosa] C85
    `--S. occidentatus (Giordani Soika 1962) [=Pseudepipona (Syneuodynerus) occidentata] C85
  Nortozumia amaliae GS77
  Hypodynerus CC85
  Cephalochilus CC85
  Ctenochilus CC85
  Raphiglossoides CC85
  Alastor CC85
    |--A. angulicollis B06
    |--A. anomalus B06
    |--A. argentinus B06
    |--A. clypeatus Brèthes 1906 B06
    |--A. elongatus B06
    |--A. nitidus Brèthes 1906 B06
    |--A. persimilis B06
    |--‘Nysson’ rottensis Meunier 1915 BM76
    `--A. schrottkyi B06
  Paramischocyttarus buyssoni CC85
  Micreumenes CC85
  Synagris Latreille 1802 CC85, L02
    |--*S. (Synagris) cornuta (Linnaeus 1758) L02, BM76 [=Vespa cornuta BM76]
    |--S. (Paragris) abyssinica Guérin in Lefebvre 1848 GS61a
    `--S. sicheliana R13
  Hypancistrocerus CC85
  Paragymnomerus CC85
  Plagiolabra Schulthess-Rechberg 1903 CC85, B06 [incl. Leontiniella Brèthes 1903 B05]
    |--P. andina Brèthes 1906 B06
    `--P. nigra [incl. Leontiniella argentina] B06
  Raphiglossa zethoides CC85, R13
  Pseudopterocheilus CC85
  Antepipona CC85
  Eustenancistrocerus CC85
  Chlorodynerus CC85
  Allodynerus CC85
    |--A. delphinalis [=Lionotus delphinalis] OK99
    |--A. mandschuricus S94
    `--A. rossi OK99
  Anterhynchium CC85
    |--A. flavomarginatum S94
    |    |--A. f. flavomarginatum S94
    |    `--A. f. micado S94
    |--A. histrionicum OK99
    `--A. melanopterum S94
  Knemodynerus CC85
  Parazumia CC85
  Psiliglossa CC85
    |--P. algeriensis CC85
    `--P. zeppelini CC85
  Pseumenes CC85
    |--P. depressus (Saussure 1856) CC85, L41 (see below for synonymy)
    |--P. imperatrix (Smith 1857) [=Eumenes imperatrix, Pareumenes (Pseumenes) imperatrix] L41
    `--P. quadrispinosus (Saussure 1856) (see below for synonymy) L41
  Tricarinodynerus guerinii CC85, GS61a
    |--T. g. guerinii GS61a
    |--T. g. deserticola Giordani Soika 1961 GS61a
    |--T. g. neptunus GS61a
    `--T. g. rubens GS61a
  Hemipterochilus CC85
  Pseudochilus CC85
  Parachilus CC85
  Pteromenes CC85
  Pseudomicrodynerus CC85
  Discoelius B05
  Labus De Saussure 1867 B06
    |--L. floricola B06
    |--L. humbertianus B06
    `--L. spiniger B06
  Oreumenes decoratus S94
  Pararrhynchium S94
    |--P. ishigakiense S94
    `--P. tsunekii Tano & Yamane 1983 I92
  Proepipona falcatum OK99
  Pareumenes L41
    |--P. acutus Liu 1941 L41
    |--P. chinensis Liu 1941 L41
    |--P. conjunctus Liu 1941 L41
    |--P. interruptus Liu 1941 L41
    |--P. obtusus Liu 1941 L41
    |--P. paeneplanus [=Eumenes paeneplanus] L41
    `--P. transitorus Liu 1941 L41
  Acarepipona pervigilans OK99
  Gribodia confluenta OK99

*Acarozumia amaliae (Saussure 1869) [=Nortonia amaliae, Montezumia (*Acarozumia) amaliae; incl. M. australensis Perkins 1908] C85

Bidentodynerus bicolor aurantiopicta (Giordani Soika 1941) C85 [=Odynerus (Rhynchium) bicolor aurantiopictus C85, Pseudepipona bicolor aurantiopicta GS61b]

Bidentodynerus bicolor flavescentulus (Giordani Soika 1941) C85 [=Odynerus (Rhynchium) bicolor flavescentulus C85, Pseudepipona bicolor flavescentula GS61b]

Bidentodynerus bicolor nigrocinctoides (Giordani Soika 1941) C85 [=Odynerus (Rhynchium) bicolor nigrocinctoides C85, Pseudepipona bicolor nigrocinctoides GS61b]

Ectopioglossa polita australensis (Meade-Waldo 1910) [=Eumenes (Pareumenes) australensis; incl. *Ec. australensis Perkins 1912 non Eumenes australensis Meade-Waldo 1910] C85

Epiodynerus nigrocinctus (Saussure 1853) [=Odynerus (Leionotus) nigrocinctus, Pseudepipona (Epiodynerus) nigrocinctus] C85

Epiodynerus tamarinus (Saussure 1853) [=Odynerus (Leionotus) tamarinus, Pseudepipona (Epiodynerus) tamarina] C85

Epiodynerus tasmaniensis (Saussure 1853) C85 [=Odynerus (Leionotus) tasmaniensis C85, Pseudepipona (Epiodynerus) tasmaniensis GS61b; incl. Rhynchium abispoides Meade-Waldo 1910 C85, Odynerus (Rhynchium) abispoides GS61b]

Leptomenoides extraneus (Saussure 1855) [=Odynerus (Leionotus) extraneus, Pachymenes extraneus; incl. O. (L.) exilis Saussure 1853 non O. exilis Schaeffer 1841] C85

Pachodynerus argentinus [=Odynerus argentinus; incl. O. argentinus var. andinus, Pachodynerus argentinus var. andinus] B06

Parifodynerus alariformis (Saussure 1856) [=Odynerus alariformis; incl. O. (Stenodyneroides) subalaris Giordani Soika 1941 non Saussure 1855] C85

Parodynerus bicincta (Fabricius 1781) [=Vespa bicincta; incl. Odynerus bizonatus Boisduval 1835, *Parodynerus bizonatus] C85

*Pseudabispa abispoides (Perkins 1912) [=Odynerus abispoides non Rhynchium abispoides Meade-Waldo 1910 (not preoc. if in dif. gen.)] C85

Pseumenes depressus (Saussure 1856) CC85, L41 [=Eumenes (Pareumenes) depressus L41, Pareumenes quadrispinosa var. depressa L41]

Pseumenes quadrispinosus (Saussure 1856) [=Eumenes (Pareumenes) quadrispinosus, Pareumenes quadrispinosa] L41

Syneuodynerus aurantiopilosellus (Giordani Soika 1962) C85 [=Pseudepipona (Syneuodynerus) aurantiopilosella C85, P. aurantiobispinosa GS61b]

*Type species of generic name indicated

REFERENCES

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

[B05] Brèthes, J. 1905. Nuevos euménidos argentinos. Anales del Museo Nacional de Buenos Aires, serie 3, 6: 21–39.

[B06] Brèthes, J. 1906. Véspidos y eumenídidos sudamericanos (nuevo suplemento). Anales del Museo Nacional de Buenos Aires, serie 3, 6: 310–377.

[C85] Cardale, J. C. 1985. Vespoidea and Sphecoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 150–303. Australian Government Publishing Service: Canberra.

[CC85] Carpenter, J. M., & J. M. Cumming. 1985. A character analysis of the North American potter wasps (Hymenoptera: Vespidae; Eumeninae). Journal of Natural History 19 (5): 877–916.

[GS61a] Giordani Soika, A. 1961a. Hymenoptera (Aculeata): Vespidae. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 440–451. Almqvist & Wiksell: Uppsala.

[GS61b] Giordani Soika, A. 1961b. Gli Odynerus sensu antiquo del continente Australiano e della Tasmania. Boll. Mus. Civ. Venezia 14: 57–202.

[GS77] Giordani Soika, A. 1977. Contributo alla conoscenza degli eumenidi australiani. Memorie della Società Entomologica Italiana 55: 109–138.

[HH07] Hines, H. M., J. H. Hunt, T. K. O'Connor, J. J. Gillespie & S. A. Cameron. 2007. Multigene phylogeny reveals eusociality evolved twice in vespid wasps. Proceedings of the National Academy of Sciences of the USA 104 (9): 3295–3299.

[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.

[L41] Liu, C. L. 1941. Revisional studies of the Vespidae of China I: the genus Pareumenes Saussure, with descriptions of six new species (Hymenoptera: Eumeninae). Notes d'Entomologie Chinoise 8 (6): 245–289.

[OK99] OConnor, B. M., & J. S. H. Klompen. 1999. Phylogenetic perspectives on mite-insect associations: the evolution of acarinaria. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 63–71. Ohio Biological Survey: Columbus (Ohio).

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[S94] Shimizu, A. 1994. Phylogeny and classification of the family Pompilidae (Hymenoptera). Tokyo Metropolitan University Bulletin of Natural History 2: 1–142.

Last updated: 19 May 2020.

Tabanomorpha

Los Angeles wormlion Vermileo, photographed by Tokuo Nakamoto.

Belongs within: Orthorrhapha.
Contains: Rhagionidae, Tabanidae.

The Tabanomorpha is a clade of flies united by the presence of a small lobe on the basal segment of the female cercus, and larvae capable of retracting their head into the thorax (Grimaldi & Engel 2005). Many members of this clade are predatory. A number of subgroups, including the Tabanidae, Athericidae and some members of the Spaniidae (previously classified with the Rhagionidae), feed on vertebrate blood (Grimaldi & Engel 2005). Other tabanomorphs include the wormlions of the genus Vermileo whose larvae construct pits like those of ant-lions that they use to capture prey. The Pelecorhynchidae are a group of moderate-sized to large flies whose predatory larvae are found in saturated soils in swamps and stream banks (Teskey 1981a). The Athericidae are medium-sized flies, often with patterned wings, with aquatic larvae mostly inhabiting rocky or gravelly areas of moving streams (Woodley 2009b). The possible stem-athericid Qiyia jurassica is known from larvae from the Middle Jurassic Daohugou Formation of Inner Mongolia that possessed adaptations for ectoparasitism such as piercing mouthparts and a ventral sucker near the front of the thorax (Chen et al. 2014).

Tabanomorpha
    |--+--Rhagionidae WT11
    |  `--Vermileonidae [Protobrachycerontidae, Vermileonoidea] WT11
    |       |--Lampromyia K09
    |       |--Protobrachyceron liasinum RJ93
    |       |--Vermitigris T81b
    |       |    |--V. fairchildi O98
    |       |    |--V. infasciatus O98
    |       |    `--V. orientalis O98
    |       `--Vermileo W09a
    |            |--V. comstocki W09a
    |            |--V. fascipennis W09a
    |            |--V. opacus T81b
    |            |--V. tibialis T81b
    |            `--V. willetti W09a
    `--+--+--Bolbomyia WT11
       |  |    |--B. macgillisi JT81
       |  |    `--B. nana WT11
       |  `--Spaniidae [Spaniinae] GE05
       |       |--Ptiolina majuscula K09, JT81
       |       |--Litoleptis alaskensis K09, JT81
       |       |--Spania nigra K09, JT81
       |       |    |--S. n. nigra JT81
       |       |    `--S. n. americana JT81
       |       |--Symphoromyia GE05
       |       |    |--S. hirta JT81
       |       |    |--S. (Axinicera) inurbana JT81
       |       |    `--S. montana M81
       |       `--Chrysopilus WT11
       |            |--C. ferruginosus O98
       |            |--C. flavibarbis K09
       |            |--C. gratiosus CM91
       |            |--C. ornatus K09
       |            |--C. proximus K09
       |            `--C. thoracicus WT11
       `--+--Batgana GE05
          |--Shulmas GE05
          `--+--Pelecorhynchidae WT11
             |    |--Bequaertomyia jonesi T81a
             |    |--Pseudoerinna K09
             |    |--Pelecorhynchus Macq. 1850 [=Caenopnyga Thoms. 1868] R00
             |    |    |--*P. maculipennis Macq. 1850 [=*Caenopnyga maculipennis] R00
             |    |    |--P. aurantiacus Ricardo 1900 R00
             |    |    |--P. darwini Ricardo 1900 R00
             |    |    |--P. eristaloides (Walker 1848) [=Silvius eristaloides] R00
             |    |    |--P. fulvus CM91
             |    |    |--P. fusconiger (Walker 1848) [=Silvius fusconiger] R00
             |    |    `--P. personatus (Walker 1848) [=Silvius personatus; incl. P. ornatus Schiner 1866] R00
             |    `--Glutops K09
             |         |--G. bandus T81a
             |         |--G. rossi T81a
             |         `--G. singularis WT11
             `--+--Oreoleptis [Oreoleptidae] WT11
                |    `--O. torrenticola WT11
                `--+--Tabanidae WT11
                   `--Athericidae WT11
                        |  i. s.: Atrichops O98
                        |--Qiyia Chen, Wang et al. 2014 CW14
                        |    `--*Q. jurassica Chen, Wang et al. 2014 CW14
                        `--+--Dasyomma [Dasyommatinae] W09b
                           `--Athericinae W09b
                                |--Xeritha W09b
                                |--Suragina W09b
                                |    |--S. concinna W09b
                                |    |--S. latipennis W09b
                                |    `--S. longipes W09b
                                `--Atherix W09b
                                     |--A. ibis O98
                                     |    |--A. i. ibis O98
                                     |    `--A. i. japonica O98
                                     |--A. lantha W81
                                     |--A. maculata [=Rhagio (Atherix) maculatus] G20
                                     |--A. peringueyi S61
                                     `--A. variegata W09b

*Type species of generic name indicated

REFERENCES

[CW14] Chen, J., B. Wang, M. S. Engel, T. Wappler, E. A. Jarzembowski, H. Zhang, X. Wang, X. Zheng & J. Rust. 2014. Extreme adaptations for aquatic ectoparasitism in a Jurassic fly larva. eLife 3: e02844.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[JT81] James, M. T., & W. J. Turner. 1981. Rhagionidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 483–488. Research Branch, Agriculture Canada.

[K09] Kerr, P. H. 2009. Rhagionidae (snipe flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 485–489. NRC Research Press: Ottawa.

[M81] McAlpine, J. F. 1981. Morphology and terminology—adults. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 9–63. Research Branch, Agriculture Canada.

[O98] Oosterbroek, P. 1998. The Families of Diptera of the Malay Archipelago. Brill: Leiden.

[R00] Ricardo, G. 1900. Notes on the Pangoninae of the family Tabanidae in the British Museum collection. Annals and Magazine of Natural History, series 7, 5: 97–121, pl. 1.

[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.

[S61] Stuckenberg, R. R. 1961. Diptera (Nematocera): Thaumaleidae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 409–412. Almqvist & Wiksell: Uppsala.

[T81a] Teskey, H. J. 1981a. Pelecorhynchidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 459–461. Research Branch, Agriculture Canada.

[T81b] Teskey, H. J. 1981b. Vermileonidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 529–532. Research Branch, Agriculture Canada.

[W81] Webb, D. W. 1981. Athericidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 479–482. Research Branch, Agriculture Canada.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

[W09a] Woodley, N. E. 2009a. Vermileonidae (worm-lions). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 481–483. NRC Research Press: Ottawa.

[W09b] Woodley, N. E. 2009b. Athericidae (athericid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 491–493. NRC Research Press: Ottawa.

Last updated: 12 April 2022.