Caloneurodea

Reconstruction of Paleuthygramma tenuicornis, from Grimaldi & Engel (2005).

Belongs within: Archaeorthoptera.

The Caloneurodea are an extinct group of cursorial insects known from the Late Carboniferous and Permian (Grimaldi & Engel 2005). They are little known with most species represented by few specimens.

Characters (from Grimaldi & Engel 2005): Antennae long, multisegmented; fore and hind wings with similar shape, venation and texture; wing veins strongly convex and concave, CuA and CuP unbranched and nearly parallel (occasionally fused) in both wing pairs; hind wing with anal area absent; tarsi five-segmented; cerci (where known) unsegmented.

Caloneurodea [Caloneurida]
    |  i. s.: Sypharoptera R02
    |         Sthenarocera R02
    |         Eohymen R02
    |         Tshecalculus R02
    |         Paoliola gourlei R02
    |         Geroneura wilsoni R02
    |         Boltonaloneura Rasnitsyn 2002 R02
    |           `--*B. subtilis (Bolton 1925) [=Caloneura subtilis] R02
    |         Aspidoneura flexa K-P91
    |--Pseudobiella [Permobiellidae] R02
    |--Anomalogrammatidae R02
    |--Amboneuridae R02
    |--Plesiogrammatidae R02
    |--Protokollaria R02 [Protokollariidae RJ93]
    |--Caloneura [Caloneuridae] R02
    |    `--*C. dawnsoni Brongniart 1885 BN02
    |--Euthygrammatidae [Gelastopteridae] R02
    |    |--Gelastopteron gracile R02
    |    `--Euthygramma parallelum Martynov 1938 BN02
    |--Paleuthygramma [Paleuthygrammatidae] GE05
    |    |--P. acutum R02
    |    |--P. tenuicornis GE05
    |    `--P. tenuis K-P91
    `--Apsidoneuridae R02
         |--Homaloptila similis (Meunier 1911) R02, BN02
         `--Apsidoneura R02
              |--A. flexa RJ93
              `--A. sottyi Burnham 1984 RJ93

*Type species of generic name indicated

REFERENCES

[BN02] Béthoux, O., & A. Nel. 2002. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1–88.

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

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[R02] Rasnitsyn, A. P. 2002. Cohors Cimiciformes Laicharting, 1781. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 104–115. 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.

Last updated: 25 May 2022.

Meliponini

Trigona carbonaria, photographed by Peter O.

Belongs within: Corbiculata.

The Meliponini, stingless bees, are a diverse group of bees in the tropics. As well as the reduced sting, they are characterised by a weakened distal forewing venation (Engel 2001).

Characters (from Engel 2001): Minute to moderate size (ca. 1.5–13 mm long), sparsely to moderately pubescent bees. Mandible without outer mandibular grooves. Labral width three to four times length. Clypeus variously produced, typically gently convex and not protuberant in profile. Compound eyes typically bare. Supra-alar carina absent; scutellum broadly rounded posteriorly and variously produced (i.e., ranging from projecting over metanotum and propodeum to not projecting at all). Claws of female simple; arolium strong and present; metatibial spurs absent; malus of strigilis without anterior velum; metabasitarsus without auricle; metatibia with penicillum. Distal venation of forewing weakened; marginal cell apex typically open; pterostigma present, moderate to large in size, much longer than prestigma, r-rs arising near midpoint, margin within marginal cell convex; 1m-cu, when present, angled; hind wing with distinct jugal lobe, lobe broadly and deeply incised; hamuli reduced; wing membrane without alar papillae. Sting reduced.

<==Meliponini [Lestrimelittini, Meliponites, Trigonini]
    |  i. s.: Plebeia CSD10
    |         Tetragonisca angustula TM03
    |         Partamona cupeira B04
    |         Cretotrigona Engel 2000 E01
    |           `--*C. prisca (Michener & Grimaldi 1988) [=Trigona (Trigona) prisca] E01
    |         Meliponorytes Tosi 1896 E01
    |           |--*M. succini Tosi 1896 E01
    |           `--M. sicula E01
    |         Proplebeia Michener 1982 E01
    |           |--*P. dominicana (Wille & Chandler 1964) (see below for synonymy) E01
    |           |--P. tantilla E01
    |           `--P. vetusta E01
    |         Dactylurina CSD10
    |--+--+--Tetragonula carbonaria (Smith 1854) CSD10
    |  |  `--+--+--Hypotrigona gribodoi (Magretti 1884) CSD10
    |  |     |  `--Kelneriapis Sakagami 1978 CSD10, E01 [=Kelnermelia Moure in Moure & Camargo 1978 E01]
    |  |     |       `--*K. eocenica (Kelner-Pillault 1969) (see below for synonymy) E01
    |  |     `--+--Liotrigonopsis Engel 2001 CSD10, E01
    |  |        |    `--*L. rozeni Engel 2001 E01
    |  |        `--Liotrigona CSD10
    |  |             |--L mahafalya E01
    |  |             `--L. vetula E01
    |  `--+--Plebeina hildebrandti (Friese 1900) CSD10
    |     `---+--Axestotrigona ferruginea (Lepeletier 1836) CSD10
    |         `--Meliponula CSD10
    |              |--M. bocandei CSD10
    |              `--M. erythra E01
    `--+--Melipona Illiger 1806 CSD10, E01
       |    |--M. alinderi BB01
       |    |--M. domestica D59
       |    |--M. quadrifasciata TM03
       |    `--M. rufiventris TM03
       `--+--Lestrimelitta Friese 1903 CSD10, E01
          `--+--Scaura latitarsis (Friese 1900) CSD10
             `--+--Scaptotrigona hellwegeri (Friese 1900) CSD10
                `--+--Cephalotrigona capitata (Smith 1854) CSD10
                   `--Trigona Jurine 1807 CSD10, E01
                        |--T. amalthea B04
                        |    |--T. a. amalthea B04
                        |    `--T. a. silvestriana B04
                        |--T. carbonaria E01
                        |--T. fulviventris TM03
                        |--T. fuscipennis Friese 1900 CSD10
                        |--T. hyalinata B04
                        |    |--T. h. hyalinata B04
                        |    `--T. h. amazonensis B04
                        |--T. iridipennis [incl. Meliponorytes devictus Cockerell 1921] E01
                        |--T. laeviceps Z02
                        `--T. lurida E01

*Kelneriapis eocenica (Kelner-Pillault 1969) [=Hypotrigona eocenica, *Kelnermelia eocenica, Tetragonula (*Kelneriapis) eocenica, Trigona (Hypotrigona) eocenica] E01

*Proplebeia dominicana (Wille & Chandler 1964) [=Trigona (Liotrigona) dominicana, T. (*Proplebeia) dominicana] E01

*Type species of generic name indicated

REFERENCES

[BB01] Bowestead, S., R. G. Booth, A. Silpinski & J. F. Lawrence. 2001. The genus Cleidostethus Arrow, 1929 reappraisal and transfer from Coccinellidae to Corylophidae (Coleoptera: Cucujoidea). Annales Zoologici 51 (3): 319-323.

[B04] Brown, B. V. 2004. Revision of the subgenus Udamochiras of Melaloncha bee-killing flies (Diptera: Phoridae: Metopininae). Zoological Journal of the Linnean Society 140: 1-42.

[CSD10] Cardinal, S., J. Straka & B. N. Danforth. 2010. Comprehensive phylogeny of apid bees reveals the evolutionary origins and antiquity of cleptoparasitism. Proceedings of the National Academy of Sciences of the USA 107 (37): 16207-16211.

[D59] Darwin, C. 1859. The Origin of Species by Means of Natural Selection, or the preservation of favoured races in the struggle for life 1^st ed. John Murray: London. (reprinted 1967. Atheneum: New York; 1968. Penguin Books: London)

[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.

[TM03] Teixeira, A. C. P., M. M. Marini, J. R. Nicoli, Y. Antonini, R. P. Martins, M.-A. Lachance & C. A. Rosa. 2003. Starmerella meliponinorum sp. nov., a novel ascomycetous yeast species associated with stingless bees. International Journal of Systematic and Evolutionary Microbiology 53: 339-343.

[Z02] Zherikhin, V. V. 2002. Pattern of insect burial and conservation. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 17-63. Kluwer Academic Publishers: Dordrecht.

Electrapini

Specimen of Protobombus indecisus in Baltic amber, from Engel (2001).

Belongs within: Corbiculata.

The Electrapini are an extinct group of bees, similar to modern bumblebees, known from the Eocene of Europe.

Characters (from Engel 2001): Moderately sized (ca. 5.5–16 mm long), robust, densely pubescent bees. Mandible with weakened outer mandibular grooves. Labral width 2–3 times length. Clypeus slightly convex and weakly protuberant in lateral view or flat (in Thaumastobombus). Compound eyes bare or with sparse, microscopic setae. Supra-alar carina present; scutellum broadly rounded posteriorly and variously produced (ranging from projecting over metanotum and propodeum to over metanotum only). Claws of female with inner tooth; arolium strong and present; single, reduced metatibial spur present; malus of strigilis with short, thickened anterior prong in addition to primary ventral velum; metabasitarsus with distinct auricle at base; metatibia without penicillum. Distal venation of forewing strong and present; marginal cell large and narrowly rounded at apex, slightly offset from wing margin, truncate or feebly appendiculate, cell longer than distance from its apex to wing apex; pterostigma present and short to moderately sized, much longer than prestigma, r-rs arising near to just after midpoint, margin within marginal cell variously produced; 1m-cu relatively straight; hind wing with distinct jugal lobe, lobe broadly and deeply incised; hamuli numerous or reduced (numerous in Electrapis and Protobombus, reduced in Thaumastobombus); wing membrane without alar papillae. Sting not reduced; with or without microscopic barbs.

<==Electrapini [Electrapina]
    |--Thaumastobombus Engel 2001 E01
    |    `--*T. andreniformis Engel 2001 E01
    |--Electrapis Cockerell 1908 WE03 [=Elektrapis (l. c.) E01; incl. Eckfeldapis Lutz 1993 E01]
    |    |--*E. meliponoides (Buttel-Reepen 1906) [=Apis meliponoides] E01
    |    |--E. electrapoides (Lutz 1993) WE03 [=*Eckfeldapis electrapoides E01]
    |    |--E. krishnorum Engel 2001 E01
    |    |--E. martialis (Cockerell 1908) [=Chalcobombus martialis] E01
    |    |--E. micheneri Wappler & Engel 2003 WE03
    |    |--E. prolata Engel & Wappler in Wappler & Engel 2003 WE03
    |    `--E. tornquisti Cockerell 1908 (see below for synonymy) E01
    `--Protobombus Cockerell 1908 WE03 [incl. Chalcobombus Cockerell 1908 E01, Sophrobombus Cockerell 1908 E01]
         |--*P. indecisus Cockerell 1908 (see below for synonymy) E01
         |--P. basilaris Engel 2001 E01
         |--P. fatalis (Cockerell 1908) [=*Sophrobombus fatalis] E01
         |--P. hirsutus (Cockerell 1908) [=Chalcobombus hirsutus] E01
         |--P. messelensis Engel & Wappler in Wappler & Engel 2003 WE03
         |--P. pristinus Wappler & Engel 2003 WE03
         `--P. tristellus Cockerell 1909 [=Electrapis (Protobombus) tristellus] E01

Electrapis tornquisti Cockerell 1908 [incl. E. bombusoides Kelner-Pillault 1974, E. cockerelli Bischoff in Kelner-Pillault 1974 (n. n.), E. cokerelli (l. c.), E. hermenaui Bischoff in Kelner-Pillault 1974 (n. n.)] E01

*Protobombus indecisus Cockerell 1908 [=Electrapis (Protobombus) indecisus; incl. *Chalcobombus humilis Cockerell 1908, Electrapis (Electrapis) apoides Manning 1960] E01

*Type species of generic name indicated

REFERENCES

[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.

[WE03] Wappler, T., & M. S. Engel. 2003. The Middle Eocene bee faunas of Eckfeld and Messel, Germany (Hymenoptera: Apoidea). Journal of Paleontology 77 (5): 908-921.

Apidae

Centris analis, photographed by Stanislav Krejčík.

Belongs within: Apiformes.
Contains: Anthrena, Anthophorini, Osirini, Nomadinae, Melectini, Corbiculata, Xylocopinae, Eucerini.

The Apidae are a diverse group of long-tongued bees in which the clypeus is bent posteriorly lateral to the narrowed labral articulation (Engel 2001). The apids include both solitary and eusocial taxa, with the eusocial taxa all contained within the Corbiculata. A large clade including the Nomadinae, Osirini and Melectini is cleptoparasitic, laying their eggs in the brood cells of other bees. The genus Ctenoplectrina is also cleptoparasitic, but belongs within the Ctenoplectrini which includes a group of bees specialised for collecting floral oils, pollen and nectar from Cucurbitaceae. The Centridini are also often collectors of floral oils but with their preferred host plants belonging to the Malpighiaceae.

Characters (from Engel 2001): Single subantennal suture; suture meeting lower margin of antennal socket. Facial foveae absent. Lower lateral margins of clypeus bent posteriorly on either side of labrum. Flabellum present; glossa acute; labial palpus with first two segments flattened, sheath-like, and elongate; submentum strongly V-shaped and sclerotized, submental arms articulating with cardines slightly above cardo-stipital articulations. Galeal comb absent; stipital comb and concavity present; basistiptial process elongate. Pre-episternal groove absent. Mesocoxa entirely exposed. Metabasitibial frequently present. Jugal lobe short. Strong metasomal scopa absent; scopa variously formed on metafemur and metatibia, metatibia sometimes developed into a corbicula (in non-parasitic females). Metapostnotum setose. Pygidial plate and fimbria of female typically present.

<==Apidae [Anthophoridae, Anthophorinae, Apiariae, Apinae, Apiti, Euceriti, Parasitae, Tetrapediini, Xylocopiformes]
    |  i. s.: Pygomelissa Engel & Wappler in Wappler & Engel 2003 WE03
    |           `--*P. lutetia Engel & Wappler in Wappler & Engel 2003 WE03
    |         Halictoides inermis MS01
    |         Rhophites quinquespinosus MS01
    |         Anthrena MS01
    |         Entechnia taurea [=Melitoma taurea] R35
    |         Sarapoda R35
    |           |--S. bimaculata R35
    |           `--S. bombiformis C08
    |--+--Anthophorini CSD10
    |  `--+--Osirini CSD10
    |     `--+--Nomadinae CSD10
    |        `--Melectini CSD10
    `--+--Centridini CSD10
       |    |--Epicharis analis Lepeletier 1841 CSD10
       |    `--+--Corbiculata CSD10
       |       `--Centris CSD10
       |            |  i. s.: C. cinerea R35
       |            |         C. hoplopoda GEEL09
       |            |         C. murales R35
       |            |         C. pallida FJ01
       |            |--+--C. analis (Fabricius 1804) CSD10
       |            |  `--C. longimana Fabricius 1804 CSD10
       |            `--+--C. dimidiata (Olivier 1789) CSD10
       |               `--+--C. hoffmanseggiae Cockerell 1897 CSD10
       |                  `--+--C. atripes Mocsáry 1899 CSD10
       |                     `--C. decolorata Lepeletier 1841 CSD10
       `--+--+--Xylocopinae CSD10
          |  `--Tetrapedia CSD10
          |      |--T. (Tetrapedia) diversipes CSD10
          |      `--T. maura Cresson 1878 CSD10
          `--+--Ctenoplectrini [Ctenoplectridae, Ctenoplectrinae] CSD10
             |    |--Ctenoplectrina Cockerell 1930 E01
             |    `--Ctenoplectra Kirby in Kirby & Spence 1826 E01
             |         |--*C. chalybea Smith 1858 M65
             |         |--C. albolimbata Magretti 1895 CSD10
             |         |--C. australica Cockerell 1926 M65
             |         `--C. bequaerti Cockerell 1930 CSD10
             `--+--+--Ancyloscelis chilensis CSD10, R35
                |  `--Exomalopsini [Exomalopsinae] CSD10
                |       |--Anthophorula completa (Cockerell 1935) CSD10
                |       `--Exomalopsis herbsti R35
                `--+--+--Eucerini CSD10
                   |  |--Teratognathini E01
                   |  `--Ancyla CSD10
                   |       |--A. anatolica Warncke 1979 CSD10
                   |       `--A. asiatica Friese 1922 CSD10
                   `--+--Tapinotaspidini CSD10
                      |    |--+--Tapinotaspoides CSD10
                      |    |  `--Caenonomada CSD10
                      |    `--+--Paratetrapedia CSD10
                      |       `--Arhysoceble picta (Friese 1899) CSD10
                      `--Emphorini CSD10
                           |--Alepidosceles CSD10
                           `--+--+--Meliphilopsis CSD10
                              |  `--Diadasia bituberculata (Cresson 1878) CSD10
                              `--+--Melitoma [Melitomini] CSD10
                                 |    `--M. chilensis BM76
                                 `--+--Diadasina distincta (Holmberg 1903) CSD10
                                    `--Ptilothrix CSD10

*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.

[CSD10] Cardinal, S., J. Straka & B. N. Danforth. 2010. Comprehensive phylogeny of apid bees reveals the evolutionary origins and antiquity of cleptoparasitism. Proceedings of the National Academy of Sciences of the USA 107 (37): 16207–16211.

[C08] Cheel. 1908. Notes and exhibits. Proceedings of the Linnean Society of New South Wales 33: 287.

[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.

[FJ01] Field, L. H., & T. H. Jarman. 2001. Mating behaviour. In: Field, L. H. (ed.) The Biology of Wetas, King Crickets and Their Allies pp. 317–332. CABI Publishing: Wallingford (UK).

[GEEL09] Greathead, D. J., N. L. Evenhuis & C. J. Einicker Lamas. 2009. Bombyliidae (bee 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. 565–576. NRC Research Press: Ottawa.

[M65] Michener, C. D. 1965. A classification of the bees of the Australian and South Pacific regions. Bulletin of the American Museum of Natural History 130: 1–362.

[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.

[R35] Rayment, T. 1935. A Cluster of Bees: Sixty essays on the life-histories of Australian bees, with specific descriptions of over 100 new species. Endeavour Press: Sydney.

[WE03] Wappler, T., & M. S. Engel. 2003. The Middle Eocene bee faunas of Eckfeld and Messel, Germany (Hymenoptera: Apoidea). Journal of Paleontology 77 (5): 908–921.

Last updated: 22 June 2018.

Megachilidae

Lithurge scabrosus, copyright Aubrey Moore.

Belongs within: Apiformes.
Contains: Anthidiini, Osmiini.

The Megachilidae are a group of long-tongued bees generally characterised by a well-developed scopa (array of long hairs) on the metasomal sterna. The plesiomorphic scopa on the hind legs of other bee families is reduced in most megachilids; basal members of the family that retain a scopa-like array of setae on the hind legs have been treated as a subfamily Fideliinae but phylogenetic analysis indicates that this group is paraphyletic (Engel 2001; Cardinal et al. 2010).

Characters (from Engel 2001): Labrum broadly articulated to clypeus, frequently longer than wide. Single subantennal suture; suture meets lower margin of antennal socket in Fideliinae, meets outer margin of antennal socket in Megachilinae and Lithurginae. Facial foveae absent. Lower lateral margins of clypeus not bent posteriorly on either side of labrum. Flabellum present; glossa acute; labial palpus with first two segments flattened, sheath-like, and elongate; submentum strongly V-shaped and sclerotized, submental arms articulating with cardines slightly above cardo-stipital articulations. Galeal comb absent; stipital comb and concavity present; basistiptial process elongate. Pre-episternal groove absent. Mesocoxa entirely exposed. Metabasitibial plate variable (present in Protolithurgini and some Lithurgini along posterior margin; absent in Fideliinae and Megachilinae). Jugal lobe short. Metasomal scopa present (in nonparasitic females). Metapostnotum setose. Pygidial plate and fimbria of female present in Fideliinae and Lithurginae, absent in Megachilinae.

<==Megachilidae [Fideliidae, Fideliinae, Fideliini, Megachiliformes]
    |  i. s.: Heteranthidium C90
    |           |--H. fontemvitae Schwarz 1926 S69
    |           `--H. larreae C90
    |         Parafidelia R35
    |--Neofidelia profuga Moure & Michener 1955 CSD10
    `--+--Fidelia CSD10
       |    |--F. major (Friese 1911) CSD10
       |    `--F. villosa R35
       `--+--+--+--Pararhopalites [Pararhophitini] CSD10
          |  |  |    `--P. quadratus (Friese 1898) CSD10
          |  |  `--Dioxys [Dioxyina, Dioxyini] PK17
          |  |       |--D. chalicoda Le Peletier 1849 E12
          |  |       |--D. cincta PK17
          |  |       `--D. pomonae Cockerell 1910 CSD10
          |  `--Megachilinae [Megachiles] CSD10
          |       |  i. s.: Probombus Piton 1940 CSD10, E01
          |       |           `--*P. hirsutus Piton 1940 E01
          |       |--Anthidiini PK17
          |       `--+--Osmiini CSD10
          |          `--Ctenoplectrellini [Ctenoplectrellina] CSD10
          |               |--Friccomelissa schopowi CSD10
          |               |--Glaesosmia Engel 2001 E01
          |               |    `--*G. genalis Engel 2001 E01
          |               `--Ctenoplectrella Cockerell 1909 E01
          |                    |--*C. viridiceps Cockerell 1909 (see below for synonymy) E01
          |                    |--C. cockerelli Engel 2001 E01
          |                    `--C. grimaldii Engel 2001 E01
          `--Lithurginae E01
               |--Protolithurgus Engel 2001 [Protolithurgini] E01
               |    `--*P. ditomeus Engel 2001 E01
               `--Lithurgini [Trichothurgini] CSD10
                    |--+--Microthurge CSD10
                    |  `--Trichothurgus CSD10 [incl. Lithurgomma E01]
                    |       `--T. herbsti (Friese 1905) CSD10
                    `--Lithurge Latreille 1825 PK17, M65 [=Lithurgus Berthold 1827 M65]
                         |  i. s.: L. adamiticus (Heer 1865) E01
                         |         ‘Lithurgus’ chrysurus PK17
                         |         L. echinocacti Cockerell 1898 CSD10
                         |         ‘Lithurgus’ fuscipennis R35
                         |         L. rubricatus Smith 1853 E01
                         `--L. (Lithurge) M65
                              |--*L. (L.) cornutus (Fabricius 1787) [=Andrena cornuta, *Lithurgus cornutus] M65
                              |--L. (L.) albofimbriatus Sichel 1867 M65
                              |--L. (L.) atratiformis Cockerell 1905 M65
                              |--L. (L.) atratus Smith 1853 M65
                              |--L. (L.) bractipes Perkins & Cheesman 1928 M65
                              |--L. (L.) cognatus Smith 1868 M65
                              |--L. (L.) dentipes Smith 1853 M65
                              |--L. (L.) fortis Cockerell 1929 M65
                              |--L. (L.) froggatti Cockerell 1914 M65
                              |--L. (L.) huberi M65
                              |--L. (L.) nigerrimus Krombein 1949 M65
                              |--L. (L.) rubricatus Smith 1853 M65
                              `--L. (L.) scabrosus (Smith 1859) [=Megachile scabrosa] M65

*Ctenoplectrella viridiceps Cockerell 1909 [incl. C. dentata Salt 1931, C. splendens Kelner-Pilaut 1970] E01

*Type species of generic name indicated

REFERENCES

[CSD10] Cardinal, S., J. Straka & B. N. Danforth. 2010. Comprehensive phylogeny of apid bees reveals the evolutionary origins and antiquity of cleptoparasitism. Proceedings of the National Academy of Sciences of the USA 107 (37): 16207–16211.

[C90] Crawford, C. S. 1990. Scorpiones, Solifugae, and associated desert taxa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 421–475. John Wiley & Sones: New York.

[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.

[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.

[M65] Michener, C. D. 1965. A classification of the bees of the Australian and South Pacific regions. Bulletin of the American Museum of Natural History 130: 1–362.

[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.

[R35] Rayment, T. 1935. A Cluster of Bees: Sixty essays on the life-histories of Australian bees, with specific descriptions of over 100 new species. Endeavour Press: Sydney.

[S69] Steyskal, G. C. 1969. The mistreatment of the Latin genitive case in forming names of parasites. Systematic Zoology 18 (3): 339–342.

Last updated: 13 September 2017.

Dorylinae

Driver ants Dorylus helvolus, photographed by Alex Wild.

Belongs within: Formicidae.

The Dorylinae, the army and driver ants, are a group of ants that have only temporary nests, with the colony moving as a whole as it forages for food. The name is used here in a broad sense; many sources divide the dorylines between a number of smaller subfamilies. The Dorylinae include some of the largest ants, with females of Dorylus species being up to 5 cm in length.

See also: Ants on the move (Taxon of the Week: Dorylidae).

Characters (from Wheeler & Wheeler 1990): Eyes absent or vestigial; pedicel usually of two segments in worker (one in female and male); clypeus short; frontal carinae short and vertical, not covering antennal insertions; antennae usually short; epinotum usually unarmed.

<==Dorylinae [Dorylidae] WW90
    |  i. s.: Nomamyrmex WW90
    |--+--Aenictogiton [Aenictogitoninae] BS06
    |  `--Dorylus Fabr. 1793 BS06, B03 (see below for synonymy)
    |       |--*D. helvolus B03
    |       |--D. brevinodosa [=D. (Typhlopone) brevinodosa] E89
    |       |--D. fuscus Emery 1889 E89
    |       |--D. labiatus Shuck. 1840 (see below for synonymy) B03
    |       |--D. laevigatus (Smith 1857) (see below for synonymy) B03
    |       |--D. nigricans I92
    |       |--D. oraniensis (Lucas 1849) E89, E12 [=Typhlopona oraniensis E12, [=D. (Typhlopone) oraniensis E89]
    |       |--D. orientalis Westw. 1835 B03 (see below for synonymy) B03
    |       `--D. punctata [=D. (Typhlopone) punctata] E89
    `--+--‘Cerapachys’ sexspinus BS06
       `--+--+--Neivamyrmex nigrescens BS06 [=Eciton nigrescens S57]
          |  `--+--Cheliomyrmex morosus BS06
          |     `--Eciton [Ecitoninae] BS06
          |          |--E. burchelli MO02
          |          |--E. dulcius WP99
          |          |--E. hamatum H79
          |          `--E. vagans BS06
          `--Aenictus Shuckard 1840 BS06, B03 [incl. Typhlatta Smith 1858 B03; Aenictinae]
               |--*A. ambiguus Shuckard 1840 B03
               |--A. aitkeni Forel 1901 B03
               |--A. aratus Forel 1900 TB85
               |--A. arya Forel 1901 B03
               |--A. binghami Forel 1901 B03
               |--A. brevicornis (Mayr 1878) [=Typhlatta brevicornis] B03
               |--A. certus Westwood 1842 B03
               |--A. ceylonicus (Mayr 1866) TB85 (see below for synonymy)
               |--A. clavatus Forel 1901 [incl. A. clavatus var. kanarensis] B03
               |--A. clavitibia Forel 1901 B03
               |--A. cornutus N-ZLI11
               |--A. dentatus N-ZLI11
               |--A. eugenii BS06
               |--A. feae Emery 1889 E89
               |--A. fergusoni Forel 1901 (see below for synonymy) B03
               |--A. glabratus N-ZLI11
               |--A. gleadowi Forel 1901 B03
               |--A. grandis Bingham 1903 B03
               |--A. greeni Bingham 1903 B03
               |--A. hilli Clark 1928 TB85
               |--A. laeviceps (Smith 1858) [=Typhlatta laeviceps] B03
               |--A. latiscapus Forel 1901 B03
               |--A. longi Forel 1901 B03
               |--A. martini Forel 1901 B03
               |--A. pachycerus (Smith 1858) (see below for synonymy) B03
               |--A. peguensis Emery 1894 B03
               |--A. philiporum Wilson 1964 TB85
               |--A. pubescens Smith 1859 B03
               |--A. punensis Forel 1901 B03
               |--A. shuckardi Forel 1901 B03
               |--A. turneri Forel 1900 H13, TB85
               |--A. westwoodi Forel 1901 B03
               `--A. wroughtoni Forel 1890 B03

Aenictus ceylonicus (Mayr 1866) TB85 [=Typhlatta ceylonica TB85; incl. A. deuqueti Crawley 1923 TB85, A. exiguus Clark 1934 TB85, A. ceylonicus var. latro B03]

Aenictus fergusoni Forel 1901 [incl. A. fergusoni var. hodgsoni, A. fergusoni var. montanus, A. fergusoni var. piltzi] B03

Aenictus pachycerus (Smith 1858) [=Eciton pachycerus; incl. Typhlatta bengalensis Mayr 1878, Aenictus bengalensis] B03

Dorylus Fabr. 1793 BS06, B03 [incl. Alaopone Emery 1881 B03, Dichthadia Gerst. 1863 B03, Typhlopone Westwood 1840 B03]

Dorylus labiatus Shuck. 1840 [=D. fulvus var. labiatus; incl. D. hindostanus Smith 1859, D. (Typhlopone) laeviceps] B03

Dorylus laevigatus (Smith 1857) [=Typhlopone laevigata; incl. Do. (Typhlopone) breviceps Emery 1889, Dichthadia glaberrima Gerst. 1863, Do. klugi Emery 1887] B03

Dorylus orientalis Westw. 1835 B03 [incl. Labidus (Typhlopone) curtisi Shuck. 1840 B03, D. fuscus Emery 1889 B03, E89, D. longicornis Shuck. 1840 B03, Alaopone oberthuri Emery 1881 B03]

*Type species of generic name indicated

REFERENCES

[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.

[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.

[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 2a 7: 485–520.

[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.

[H13] Heterick, B. E. 2013. A taxonomic overview and key to the ants of Barrow Island, Western Australia. Records of the Western Australian Museum Supplement 83: 375–404.

[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.

[I92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[MO02] Machado, G., & P. S. Oliveira. 2002. Maternal care in the Neotropical harvestman Bourguyia albiornata (Arachnida: Opiliones): oviposition site selection and egg protection. Behaviour 139: 1509–1524.

[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.

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

[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.

Monomorium

Monomorium longipes, copyright Zach Lieberman/antweb.org.

Belongs within: Myrmicinae.

Monomorium is a cosmopolitan genus of small ants, most diverse in the Old World tropics with relatively few species native to the Americas. Colonies are often polygynous and may possess either winged or ergatoid queens or a combination of both. Despite their small size, a number of species of Monomorium are significant pests. These include the pharaoh ant Monomorium pharaonis and the Singapore ant M. destructor. The latter species is known to cause damage to infrastructure by chewing through cabling (Heterick 2009). Monophyly of the genus has been questioned by recent studies leading to proposals to divide it between several genera.

Characters (from Wheeler & Wheeler 1990): Antenna with twelve segments; epinotum unarmed, basal face at same level as dorsum of mesonotum; postpetiol attached to anterior end of first gastric somite; gaster about equally convex above and below.

<==Monomorium Mayr 1855 FT08 (see below for synonymy)
    |--M. monomorium (Mayr 1855) Z01 [incl. Myrmica (Monomorium) carbonaria Smith 1858 B03, *Mo. minutum Mayr 1855 Z01]
    |--M. aberrans Forel 1902 B03
    |--M. aithoderum H09
    |--M. antarcticum WL04 [=Chelaner antarcticus M83]
    |--M. anthracinum H09
    |--M. aper Emery 1914 [incl. M. aper var. dubia Emery 1914] E14
    |--M. arenarium H09
    |--M. armstrongi McAreavey 1949 [=M. (Holcomyrmex) armstrongi, Chelaner armstrongi] TB85
    |--M. atomus Forel 1902 [incl. M. atomus var. integrius] B03
    |--M. australicum Forel 1907 TB85 [=M. subcoecum australicum TB85; incl. M. talpa FT08]
    |--M. barbatulum E89
    |--M. bicorne Forel 1907 H09, TB85 [=Chelaner bicornis TB85; incl. M. macareaveyi H09]
    |--M. bihamatum H09
    |--M. brachythrix H09
    |--M. broomense Forel 1915 [=M. (Mitara) laeve broomense] TB85
    |--M. centrale Forel 1910 H09, TB85 [=Chelaner centralis TB85]
    |--M. chinense Z01
    |--M. concolor Zhou 2001 Z01
    |--‘Holcomyrmex’ criniceps Mayr 1878 [incl. H. crinicipitoscabriceps Forel 1902] B03
    |--M. crinitum H09
    |--M. croceiventre Emery 1914 E14
    |--M. decuria H09
    |--M. destructor (Jerdon 1851) FT08 (see below for synonymy)
    |--M. dichroum Forel 1902 B03
    |--M. disetigerum H09
    |--M. donisthorpei Crawley 1915 [=M. (Mitara) donisthorpei] TB85
    |--M. durokoppinense H09
    |--M. edentatum Emery 1897 E14
    |--M. elegantulum H09
    |--M. eremophilum H09
    |--M. ergatogyna BS06
    |--M. euryodon H09
    |--M. falcatum (McAreavey 1949) H09, TB85 [=*Schizopelta falcata H09, Chelaner falcatus TB85]
    |--M. fieldi Forel 1910 TB85 [=M. (Martia) fieldi TB85; incl. M. (Mitara) laeve nigrius Forel 1915 TB85, H09, TB85]
    |--M. flavipes Clark 1938 [=M. (Notomyrmex) flavipes, Chelaner flavipes] TB85
    |--M. flavonigrum H09
    |--M. floricola (Jerdon 1851) FT08 (see below for synonymy)
    |--M. forcipatum Emery 1914 [=M. (*Chelaner) forcipatum, *Chelaner forcipatus] TB85
    |--M. foreli Viehmeyer 1913 [=M. (Holcomyrmex) foreli, Chelaner foreli] TB85
    |--M. fossulatum Emery 1894 B03
    |--M. fraterculus Santschi 1919 [=M. (Mitara) laeve fraterculus] TB85
    |    |--M. f. fraterculus TB85
    |    `--M. f. barretti Santschi 1928 [=M. (Lampromyrmex) fraterculus barretti] TB85
    |--M. gilberti Forel 1902 [=Chelaner gilberti] TB85
    |    |--M. g. gilberti TB85
    |    `--M. g. mediorubra Forel 1915 [=Chelaner gilberti mediorubrus] TB85
    |--‘Holcomyrmex’ glaber André 1883 [incl. H. glaber var. clarus Forel 1902, H. glabrocriniceps Forel 1902] B03
    |--M. glyciphilum (Smith 1858) [=Myrmica glyciphila] B03
    |--M. gracillimum (Smith 1861) [=Myrmica gracillima] B03
    |--M. hildebrandti H09
    |--M. howense Wheeler 1927 [=M. (Notomyrmex) howense, Chelaner howensis] TB85
    |--M. ilia Forel 1907 [=M. (Martia) ilia] TB85
    |    |--M. i. ilia TB85
    |    `--M. i. lamingtonense Forel 1915 [=M. (Mitara) ilia lamingtonensis] TB85
    |--M. indicum Forel 1902 [=M. salomonis indicum] B03
    |--M. insolescens Wheeler 1934 H13, TB85 [=M. (Notomyrmex) insolescens TB85, Chelaner insolescens TB85]
    |--M. insularis Clark 1938 [=M. (Notomyrmex) insularis, Chelaner insularis] TB85
    |--M. integrum E14
    |--M. intrudens Smith 1874 [incl. lampromyrmex gracilum Mayr 1868, M. nipponense Wheeler 1906] Z01
    |--M. kiliani Forel 1902 H09, TB85 [=Chelaner kiliani TB85]
    |    |--M. k. kiliani TB85
    |    |--M. k. obscurella Viehmeyer 1925 [=Chelaner kiliani obscurellus] TB85
    |    `--M. k. tambourinensis Forel 1915 [=Chelaner kiliani tambourinensis] TB85
    |--M. lacunosum H09
    |--M. laeve Mayr 1876 [=M. (*Mitara) laeve] TB85
    |--M. latinode Mayr 1872 FT08
    |--M. leae Forel 1913 H09, TB85 [=Chelaner leae TB85; incl. M. (Notomyrmex) hemiphaeum Clark 1934 TB85]
    |--M. legulus H09
    |--M. longi Forel 1902 B03
    |--M. longiceps Wheeler 1934 H09, TB85 [=M. (Notomyrmex) longiceps TB85, Chelaner longiceps TB85]
    |--M. longinode H09
    |--M. longipes Emery 1914 [=M. (Chelaner) longipes] E14
    |--M. luteum G89
    |--‘Chelaner’ macareaveyi Ettershank 1966 (see below for synonymy) TB85
    |--M. majeri H09
    |--M. mayri Forel 1902 Z01 (see below for synonymy)
    |--M. megalops H09
    |--M. melleum Emery 1914 E14
    |--M. micron Crawley 1925 TB85
    |--M. micula H09
    |--‘Holcomyrmex’ muticus Emery 1887 B03
    |--M. nanum H09
    |--M. nitidum E14
    |--M. occidaneus Crawley 1922 [=Chelaner occidaneus] TB85
    |--M. orientale Mayr 1879 FT08
    |--M. pergandei [=Epoecus pergandei] WW90
    |--M. pharaonis (Linnaeus 1758) FT08 (see below for synonymy)
    |--M. pubescens H09
    |--M. punctipectoris Zhou 2001 Z01
    |--M. punctulatum MC13
    |--M. rothsteini Forel 1902 H09, TB85 [=Chelaner rothsteini H09, M. (*Protholcomyrmex) rothsteini TB85]
    |    |--M. r. rothsteini TB85
    |    |--M. r. doddi Santschi 1919 [=M. (Paraholcomyrmex) rothsteini doddi, Chelaner rothsteini doddi] TB85
    |    |--M. r. humilior Forel 1910 [=Chelaner rothsteini humilior] TB85
    |    |--M. r. leda Forel 1915 [=Chelaner rothsteini leda] TB85
    |    |--M. r. squamigena Viehmeyer 1925 [=Chelaner rothsteini squamigena] TB85
    |    `--M. r. tostum Wheeler 1915 [=Chelaner rothsteini tostum] TB85
    |--M. rubriceps Mayr 1876 H13, TB85 [=Chelaner rubriceps TB85]
    |    |--M. r. rubriceps TB85
    |    |--M. r. cinctum Wheeler 1917 [=Chelaner rubriceps cinctus] TB85
    |    |--M. r. extreminigrum Forel 1915 [=Monomorium rubriceps extreminigrus] TB85
    |    `--M. r. rubra Forel 1915 [=Chelaner rubriceps rubrus] TB85
    |--M. rufiventris [=Vollenhovia rufiventris] E14
    |--M. rufonigrum H09
    |--M. sagei Forel 1902 B03
    |--M. sanguinolentum Wheeler 1927 [=M. (Notomyrmex) sanguinolentum, Chelaner sanguinolentus] TB85
    |--‘*Holcomyrmex’ scabriceps Mayr 1878 B03
    |--M. sculpturatum Clark 1934 H09, TB85 [=M. (Notomyrmex) sculpturatum TB85, Chelaner sculpturatus TB85]
    |--M. silaceum H09
    |--M. sordidum Forel 1902 H09, TB85 [=Chelaner sordidus H09]
    |    |--M. s. sordidum TB85
    |    `--M. s. nigriventris Forel 1910 [=Chelaner sordidus nigriventris] TB85
    |--M. stictonotum H09
    |--M. striatifrons H09
    |--M. subapterum Wheeler 1917 [=Chelaner subapterus] TB85
    |    |--M. s. subapterum TB85
    |    `--M. s. bogischi Wheeler 1917 [=Chelaner subapterus bogischi] TB85
    |--M. subcoecum Emery 1894 FT08
    |--M. sublamellatum H09
    |--M. subopacum (Smith 1858) (see below for synonymy) Z01
    |--M. sydneyense Forel 1902 TB85
    |    |--M. s. sydneyense TB85
    |    `--M. s. nigellum Emery 1914 [=M. (Mitara) sydneyense nigella] TB85
    |--M. tricolor Emery 1914 E14
    |--M. turneri (Forel 1910) E14, TB85 [=Vollenhovia turneri TB85, Chelaner turneri TB85]
    |--M. whitei Wheeler 1915 [=M. (Holcomyrmex) whitei, Chelaner whitei] TB85
    |--M. wroughtoni Forel 1902 B03
    `--M. xantheklemma H09

‘Chelaner’ macareaveyi Ettershank 1966 [=Monomorium (Holcomyrmex) niger McAreavey 1949 non Holcomyrmex criniceps nigrum Forel 1902] TB85

Monomorium Mayr 1855 FT08 [incl. Chelaner Emery 1914 Z01, Corynomyrmex Viehmeyer 1916 Z01, Epixenus Emery 1908 Z01, Epoecus Emery 1892 Z01, Equestrimessor Santschi 1919 Z01, Holcomyrmex Mayr 1878 Z01, Irenidris Donisthorpe 1943 Z01, Isolcomyrmex Santschi 1917 Z01, Lampromyrmex Mayr 1868 Z01, Mitara Emery 1913 TB85, Notomyrmex Emery 1915 Z01, Paraphacota Santschi 1919 Z01, Parholcomyrmex Emery 1915 Z01, Phacota Roger 1862 Z01, Protholcomyrmex Wheeler 1922 TB85, Schizopelta McAreavey 1949 H09, Wheeleria Forel 1905 Z01, Wheleriella Forel 1907 Z01, Xenhyboma Santschi 1919 Z01, Xeromyrmex Emery 1915 Z01]

Monomorium destructor (Jerdon 1851) FT08 [=Atta destructor B03; incl. Mo. basale Mayr 1865 B03, Myrmica vastator Smith 1857 B03]

Monomorium floricola (Jerdon 1851) FT08 [=Atta floricola Z01, M. floreanum (l. c.) Z01; incl. M. anngusticlava Donisthorpe 1947 Z01, M. cinnabari Rober 1863 Z01, M. impressum Smith 1876 Z01, M. poecilum Roger 1863 Z01, M. specularis Mayr 1866 Z01]

Monomorium mayri Forel 1902 Z01 [=M. destructor var. mayri Z01, M. gracillimum var. mayri B03; incl. M. gracillimum var. karawajewi Wheeler 1922 Z01]

Monomorium pharaonis (Linnaeus 1758) FT08 [=Formica pharaonis Z01; incl. F. antiguensis Fabricius 1793 Z01, Myrmica contigua Smith 1858 Z01, My. domestica Schuckard 1838 Z01, Diplorhoptrum domesticum Gaskell 1877 Z01, My. fragilis Smith 1858 Z01, D. fugax Lucas 1858 Z01, Atta minuta Jerdon 1851 Z01, My. molesta Say 1836 Z01, My. unifasciata Bostock 1839 Z01]

Monomorium subopacum (Smith 1858) [=Myrmica subopaca, Mo. salomonis var. subopaca; incl. Mo. subopacum var. intermedium Santschi 1928, Mo. subopacum var. liberta Santschi 1927, Mo. mediterraneum Mayr 1861, Paraphacota cabrenae st. obscuripes Santschi 1921, P. surcoufi Santschi 1919] Z01

*Type species of generic name indicated

REFERENCES

[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.

[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.

[E89] Emery, C. 1889. Intorno ad alcune formiche della fauna palearctica. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 439–443.

[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.

[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.

[H13] Heterick, B. E. 2013. A taxonomic overview and key to the ants of Barrow Island, Western Australia. Records of the Western Australian Museum Supplement 83: 375–404.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

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

[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.

[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.

[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.

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

Last updated: 13 June 2020.

Myrmeciinae

Brown bull ant Myrmecia pyriformis, photographed by Pavel Krásenský.

Belongs within: Formicidae.

The Myrmeciinae, bulldog ants, are a group of often large ants with a powerful sting. Living myrmeciines are restricted to Australasia though fossil representatives are known from other continents. The genus Prionomyrmex is known from Baltic amber, late Eocene or early Oligocene in age, whereas Archimyrmex is known from the Eocene of North and South America (Ward & Brady 2003a, b). Members of the tribe Prionomyrmecini, including the fossil genera Prionomyrmex and Archimyrmex, together with the living Nothomyrmecia, have serrate, tightly-closing but non-overlapping mandibles, while Myrmecia species have curved mandibles with broadly-spaced teeth (Heterick 2009; Ward & Brady 2003). Members of the Myrmeciinae live in small colonies, with minimal functional differentiation between individuals. One species, Myrmecia inquilina, is a social parasite of other Myrmecia species that is not known to produce workers of its own (Heterick 2009).

Characters (from Ward & Brady 2003a): Mandibles multidentate and elongate, such that mandible length is three-quarters or more of head length. Palp formula 6, 4 (possibly reduced in Prionomyrmex janzeni). Clypeus with posteromesial protrusion between frontal carinae and antennal sockets. Eyes large, oval and conspicuously protruding from the surface of the head. First funicular segment less than, or approximately equal to, second funicular segment in length. Second funicular segment slender (more than twice as long as wide) and notably longer than the third funicular segment. Promesonotal suture flexible. Metacoxal cavities open (condition in Prionomyrmex unknown). Metapleural gland opening flanked above by carina-like flange and separated from ventral margin of metapleuron by a distance greater than the diameter of the opening (condition in Prionomyrmex unknown). Hind tibia with two apical spurs, the posterior one usually pectinate. Basitarsi of mid and hind legs with longitudinal sulcus. Tarsal claws bifurcate, with submedian tooth in addition to apical tooth. Petiole, postpetiole and abdominal segment 4 lacking tergosternal fusion (some fusion occurs anteriorly in the petiole of Nothomyrmecia). Third abdominal segment substantially smaller than fourth abdominal segment such that, when observed in profile under normal gastral distension, the height of third abdominal segment distinctly less than that of the fourth abdominal segment. Sting well developed (some reduction in Nothomyrmecia compared with Myrmecia). Alates with forewing Cu and M veins diverging at, or near, the cu-a crossvein; forewing crossvein m-cu joining M distad of divergence between M and Rs; forewing with two submarginal cells; hind wing jugal lobe present. Male with first antennal segment (scape) very short and stocky, such that scape length is about one-fifth of combined length of antennal segments 2–4; mesepisternum lacking distinct posterior oblique sulcus, at most a weak furrow present (condition in Prionomyrmex unclear); paramere divided by an oblique longitudinal suture into (i) an apical and ventromesial portion, to which the volsella is attached, and (ii) a proximal and dorsolateral portion (condition in Prionomyrmex unknown); aedeagus with posteroventral projection, armed at the apex with stout teeth or spines (condition in Prionomyrmex unknown).

<==Myrmeciinae
    |--Prionomyrmecini [Nothomyrmeciinae] WB03a
    |    |--Prionomyrmex Mayr 1868 WB03a
    |    |    |--P. janzeni WB03a
    |    |    `--P. longiceps WB03a
    |    `--+--Nothomyrmecia Clark 1934 WB03a
    |       |    `--*N. macrops Clark 1934 TB85
    |       `--Archimyrmex WB03b (see below for synonymy)
    |            |--*A. rostratus WB03b
    |            |--‘Ameghinoia’ piatnitzkyi WB03b
    |            `--‘Polanskiella’ smekali WB03b
    `--Myrmecia Fabricius 1804 (see below for synonymy) WB03a
         |--*M. gulosa (Fabricius 1775) [=Formica gulosa; incl. M. gulosa obscurior Forel 1922] TB85
         |--M. aberrans Forel 1900 [=M. (*Promyrmecia) aberrans] TB85
         |--M. acuta H09
         |--M. analis Mayr 1862 [incl. M. atriscapa Crawley 1925] TB85
         |--M. apicalis Emery 1883 E14
         |--M. arnoldi Clark 1951 TB85
         |--M. atrata Clark 1951 TB85
         |--M. auriventris Mayr 1870 [incl. M. auriventris athertonensis Forel 1915] TB85
         |--M. brevinoda Forel 1910 (see below for synonymy) TB85
         |--M. callima (Clark 1943) [=Promyrmecia callima] TB85
         |--M. cardigaster Brown 1953 [=M. cordata Clark 1951 non Latreille 1805] TB85
         |--M. celaena (Clark 1943) [=Promyrmecia celaena] TB85
         |--M. cephalotes (Clark 1943) [=Promyrmecia cephalotes] TB85
         |--M. chasei Forel 1894 [incl. M. pilosula mediorubra Forel 1910] TB85
         |--M. chrysogaster (Clark 1943) [=Promyrmecia chrysogaster] TB85
         |--M. clarki Crawley 1922 TB85
         |--M. comata Clark 1951 TB85
         |--M. croslandi MG06
         |--M. cydista (Clark 1943) [=Promyrmecia cydista] TB85
         |--M. decipians Clark 1951 TB85
         |--M. desertorum Wheeler 1915 (see below for synonymy) TB85
         |--M. dichospila Clark 1938 [=M. (Promyrmecia) dichospila] TB85
         |--M. dimidiata Clark 1951 TB85
         |--M. dispar (Clark 1951) [=Promyrmecia dispar] TB85
         |--M. dixoni (Clark 1943) [=Promyrmecia dixoni] TB85
         |--M. elegans (Clark 1943) [=Promyrmecia elegans] TB85
         |--M. erecta H09
         |--M. esuriens Fabricius 1804 [incl. M. tasmaniensis Smith 1858, M. walkeri Forel 1893] TB85
         |--M. eupoecila (Clark 1943) [=Promyrmecia eupoecila] TB85
         |--M. excavata (Clark 1951) [=Promyrmecia excavata] TB85
         |--M. exigua (Clark 1943) [=Promyrmecia exigua] TB85
         |--M. fasciata Clark 1951 TB85
         |--M. ferruginea Mayr 1876 [=M. nigriceps ferruginea] TB85
         |--M. flammicollis Brown 1953 TB85
         |--M. flavicoma Roger 1861 TB85
         |    |--M. f. flavicoma TB85
         |    `--M. f. minuscula Forel 1915 TB85
         |--M. forceps Roger 1861 [incl. M. forceps obscuriceps Viehmeyer 1924, M. singularis Clark 1951] TB85
         |--M. forficata (Fabricius 1787) (see below for synonymy) TB85
         |--M. froggatti Forel 1910 (see below for synonymy) TB85
         |--M. fucosa Clark 1934 [=M. (Promyrmecia) fucosa] TB85
         |--M. fulgida Clark 1951 TB85
         |--M. fulviculis Forel 1913 [=M. (Pristomyrmecia) fulvipes fulviculis] TB85
         |--M. fulvipes Roger 1861 (see below for synonymy) TB85
         |--M. fuscipes Clark 1951 H09, TB85
         |--M. gilberti Forel 1910 [=M. fulvipes gilberti; incl. M. (Pristomyrmecia) regina Santschi 1928] TB85
         |--M. gratiosa Clark 1951 TB85
         |--M. greavesi (Clark 1943) [=Promyrmecia greavesi] TB85
         |--M. harderi Forel 1910 [incl. Promyrmecia maloni Clark 1943, P. scabra Clark 1943] TB85
         |--M. hilli (Clark 1943) [=Promyrmecia hilli] TB85
         |--M. hirsuta Clark 1951 TB85
         |--M. infima Forel 1900 TB85
         |--M. inquilina Douglas & Brown 1959 TB85
         |--M. longinodis Clark 1951 TB85
         |--M. ludlowi Crawley 1922 H09, TB85 [=M. chasei ludlowi TB85]
         |--M. luteiforceps (Clark 1943) [=Promyrmecia luteiforceps] TB85
         |--M. mandibularis Smith 1858 (see below for synonymy) TB85
         |--M. maura Wheeler 1933 [=M. (Promyrmecia) aberrans maura] TB85
         |    |--M. m. maura TB85
         |    `--M. m. formosa Wheeler 1933 (see below for synonymy) TB85
         |--M. michaelseni Forel 1907 TB85
         |    |--M. m. michaelseni [incl. M. michaelseni perthensis Crawley 1922] TB85
         |    `--M. m. queenslandica Forel 1915 [incl. M. michaelseni overbecki Viehmeyer 1924] TB85
         |--M. midas Clark 1951 TB85
         |--M. mjobergi Forel 1915 TB85
         |--M. nigra Forel 1907 [=M. picta nigra] TB85
         |--M. nigriceps Mayr 1862 TB85
         |--M. nigriscapa Roger 1861 TB85
         |--M. nigrocincta Smith 1858 TB85
         |--M. nobilis (Clark 1943) [=Promyrmecia nobilis] TB85
         |--M. occidentalis (Clark 1943) [=Promyrmecia occidentalis] TB85
         |--M. opaca (Clark 1943) [=Promyrmecia opaca] TB85
         |--M. pavida Clark 1951 TB85
         |--M. petiolata Emery 1895 TB85
         |--M. picta Smith 1858 TB85
         |--M. picticeps Clark 1951 TB85
         |--M. piliventris Smith 1858 [incl. M. piliventris rectidens Forel 1910] TB85
         |--M. pilosula Smith 1858 [=M. (*Halmamyrmecia) pilosula; incl. Ponera ruginoda Smith 1858] TB85
         |--M. potteri (Clark 1951) [=Promyrmecia potteri] TB85
         |--M. pulchra Clark 1929 [incl. M. crassinoda Clark 1934, M. fallax Clark 1951, M. murina Clark 1951] TB85
         |--M. pyriformis Smith 1858 [incl. M. sanguinea Smith 1858] TB85
         |--M. regularis Crawley 1925 TB85
         |--M. rowlandi Forel 1910 [=M. tarsata rowlandi; incl. M. tarsata malandensis Forel 1915] TB85
         |--M. rubicunda (Clark 1943) [=Promyrmecia rubicunda] TB85
         |--M. rubripes Clark 1951 TB85
         |--M. rufinodis Smith 1858 [incl. M. gracilis Emery 1898] TB85
         |--M. rugosa Wheeler 1933 [=M. (Promyrmecia) michaelseni rugosa; incl. Promyrmecia ruginodis Clark 1943] TB85
         |--M. simillima Smith 1858 (see below for synonymy) TB85
         |--M. subfasciata Viehmeyer 1924 TB85
         |--M. suttoni Clark 1951 TB85
         |--M. swalei Crawley 1922 [=M. harderi swalei] TB85
         |--M. tarsata Smith 1858 TB85
         |--M. tepperi Emery 1898 TB85
         |--M. testaceipes (Clark 1943) [=Promyrmecia testaceipes] TB85
         |--M. urens Lowne 1865 [incl. M. pumilio Mayr 1866] TB85
         |--M. varians Mayr 1876 (see below for synonymy) TB85
         `--M. vindex Smith 1858 TB85
              |--M. v. vindex TB85
              `--M. v. basirufa Forel 1907 TB85

Archimyrmex WB03b [incl. Ameghinoia Viana & Haedo Rossi 1957 WB03b, WB03a, Polanskiella Rossi de Garcia 1983 WB03b, WB03a]

Myrmecia Fabricius 1804 [incl. Halmamyrmecia Wheeler 1922, Pristomyrmecia Emery 1911, Promyrmecia Emery 1911; Myrmeciini] WB03a

Myrmecia brevinoda Forel 1910 [=M. forficata brevinoda; incl. M. forficata eudoxia Forel 1915, M. pyriformis gigas Forel 1913] TB85

Myrmecia desertorum Wheeler 1915 [incl. M. vindex desertorum; incl. M. lutea Crawley 1922, M. princeps Clark 1951] TB85

Myrmecia forficata (Fabricius 1787) [=Formica forficata; incl. M. lucida Forel 1893, M. forficata rubra Forel 1910] TB85

Myrmecia froggatti Forel 1910 [incl. M. (Promyrmecia) aberrans sericata Wheeler 1933, M. (P.) aberrans taylori Wheeler 1933] TB85

Myrmecia fulvipes Roger 1861 [incl. M. (Promyrmecia) fulvipes barbata Wheeler 1933, M. (Pristomyrmecia) piliventris femorata Santschi 1928] TB85

Myrmecia mandibularis Smith 1858 [=M. (*Pristomyrmecia) mandibularis; incl. M. mandibularis aureorufa Forel 1910, M. (Promyrmecia) fulvipes caelatinoda Wheeler 1933, Promyrmecia laevinodis Clark 1943, M. (Pro.) mandibularis postpetiolaris Wheeler 1933] TB85

Myrmecia maura formosa Wheeler 1933 [=M. (Promyrmecia) aberrans formosa; incl. M. (P.) aberrans haematosticta Wheeler 1933] TB85

Myrmecia simillima Smith 1858 [incl. M. affinis Mayr 1862, M. crudelis Smith 1858, M. nigriventris Mayr 1862, M. paucidens Forel 1910, M. tricolor rogeri Emery 1914, M. spadicea Mayr 1862, M. tricolor Mayr 1862] TB85

Myrmecia varians Mayr 1876 [incl. Promyrmecia goudiei Clark 1943, P. marmorata Clark 1951, Myrmecia rufonigra Crawley 1921, P. shepherdi Clark 1943, P. wilsoni Clark 1943] TB85

*Type species of generic name indicated

REFERENCES

[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.

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

[MG06] Mallatt, J., & G. Giribet. 2006. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40: 772–794.

[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.

[WB03a] Ward, P. S., & S. G. Brady. 2003a. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361–386.

[WB03b] Ward, P. S., & S. G. Brady. 2003b. Corrigendum: Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 605.

Last updated: 20 May 2020.