Caloneurodea

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


Belongs within: Orthopteroidea.

The Caloneurodea are an extinct group of cursorial insects known from the Late Carboniferous and Permian (Grimaldi & Engel 2005).

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
    |         Protokollaria R02
    |         Sthenarocera R02
    |         Eohymen R02
    |         Tshecalculus R02
    |         Paoliola gourlei R02
    |         Geroneura wilsoni R02
    |         Pruvostiella R02
    |         Boltonaloneura Rasnitsyn 2002 R02
    |           `--*B. subtilis (Bolton 1925) [=Caloneura subtilis] R02
    |--Pseudobiella [Permobiellidae] R02
    |--Anomalogrammatidae R02
    |--Amboneuridae R02
    |--Plesiogrammatidae R02
    |--Caloneura [Caloneuridae] R02
    |    `--*C. dawnsoni Brongniart 1885 BN02
    |--Paleuthygramma [Paleuthygrammatidae] GE05
    |    |--P. acutum R02
    |    `--P. tenuicornis GE05
    |--Euthygrammatidae [Gelastopteridae] R02
    |    |--Gelastopteron gracile R02
    |    `--Euthygramma parallelum Martynov 1938 BN02
    `--Apsidoneuridae R02
         |--Homaloptila similis (Meunier 1911) R02, BN02
         `--Apsidoneura R02

*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.

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

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 1st 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: 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.

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, Apiariae, Apinae, Apiti, Euceriti, Tetrapediini]
    |--+--Anthophorini CSD10
    |  `--+--Osirini CSD10
    |     `--+--Nomadinae CSD10
    |        `--Melectini CSD10
    `--+--Centridini CSD10
       |    |--Epicharis analis Lepeletier 1841 CSD10
       |    `--+--Corbiculata CSD10
       |       `--Centris CSD10
       |            |  i. s.: 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 [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 CSD10
                |  `--Exomalopsini [Exomalopsinae] CSD10
                |       |--Anthophorula completa (Cockerell 1935) CSD10
                |       `--Exomalopsis CSD10
                `--+--+--Eucerini CSD10
                   |  |--Teratognathini E01
                   |  `--Ancyla CSD10
                   |       |--A. anatolica Wamcke 1979 CSD10
                   |       `--A. asiatica 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

Apidae incertae sedis:
  Pygomelissa Engel & Wappler in Wappler & Engel 2003 WE03
    `--*P. lutetia Engel & Wappler in Wappler & Engel 2003 WE03
  Podalirius Latreille 1802 L02
    |--P. acervorum [=Apis acervorum] L02
    |--P. crassipes [=Apis crassipes] L02
    |--P. hirsuta [=Andrena hirsuta] L02
    |--P. lanipes [=Apis lanipes] L02
    |--P. pilipes [=Apis pilipes] L02
    |--P. rotundata [=Apis rotundata] L02
    `--P. versicolor [=Apis versicolor] L02

*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.

[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 The Biology of Wetas, King Crickets and Their Allies (L. H. Field, ed.) pp. 317-332. CABI Publishing: Wallingford (UK).

[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.

[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.

[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.

Megachilidae

Osmia cornifrons, photographed by Beatriz Moisset.


Belongs within: Apiformes.

The Megachilidae are a group of long-tongued bees generally characterised by a well-developed scopa (array of long hairs) on the metasomal sterna, and subantennal sutures directed to the outer margins of the antennal sockets (Engel 2001).

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 [Fideliinae, Fideliini]
    |  i. s.: Heteranthidium larreae C90
    |--Neofidelia profuga Moure & Michener 1955 CSD10
    `--+--Fidelia major (Friese 1911) CSD10
       `--+--+--Pararhopalites [Pararhophitini] CSD10
          |  |    `--P. quadratus (Friese 1898) CSD10
          |  `--Dioxys [Dioxyina, Dioxyini] CSD10
          |       |--D. chalicoda Le Peletier 1849 E12
          |       `--D. pomonae Cockerell 1910 CSD10
          `--Megachilinae [Megachiles] CSD10
               |  i. s.: Chalicodoma ustulatum OC01, B88
               |         Probombus Piton 1940 CSD10, E01
               |           `--*P. hirsutus Piton 1940 E01
               |--Lithurginae E01
               |   |--Protolithurgus Engel 2001 [Protolithurgini] E01
               |   |    `--*P. ditomeus Engel 2001 E01
               |   `--Lithurgini [Trichothurgini] E01
               |        |--+--Microthurge CSD10
               |        |  `--Trichothurgus CSD10 [incl. Lithurgomma E01]
               |        |       `--T. herbsti (Friese 1905) CSD10
               |        `--Lithurgus CSD10
               |             |--L. adamiticus (Heer 1865) E01
               |             |--L. echinocacti Cockerell 1898 CSD10
               |             `--L. rubricatus Smith 1853 E01
               `--+--Anthidiini CSD10
                  |    |  i. s.: Lithanthidium Cockerell 1911 E01
                  |    |           `--*L. pertriste Cockerell 1911 E01
                  |    |         Archianthidium OC01
                  |    |--Trachusa larreae (Cockerell 1897) CSD10
                  |    `--+--Anthidium [Anthidiina] CSD10
                  |       |    |--A. klugii Lucas 1849 E12
                  |       |    `--A. porterae Cockerell 1900 CSD10
                  |       `--+--Stelis linsleyi Timberlake 1941 CSD10
                  |          `--+--Paranthidium jugatorium (Say 1824) CSD10
                  |             `--+--Anthidiellum notatum (Latreille 1809) CSD10
                  |                `--Dianthidium subparvum Swenk 1914 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
                     `--Osmiini [Chelostomidae, Glyptapidae, Osmiites, Trypetini] E01
                          |  i. s.: Ochreriades E01
                          |         Apiaria Germar 1839 E01
                          |           `--*A. dubia Germar 1839 E01
                          |         Glyptapis Cockerell 1909 [Glyptapina, Glyptapinae] E01
                          |           |--*G. mirabilis Cockerell 1909 E01
                          |           |--G. densopunctata Engel 2001 E01
                          |           |--G. disareolata Engel 2001 E01
                          |           `--G. fuscula Cockerell 1909 (see below for synonymy) E01
                          |--+--Afroheriades hyalinus CSD10
                          |  `--Megachilini [Chalicodomopsini, Coelioxyinae]
                          |       |--Coelioxys alternata Say 1837 CSD10
                          |       `--Megachile Latreille 1802 E01
                          |            |  i. s.: M. angelarum Cockerell 1902 CSD10
                          |            |         M. ericetorum Lepeletier 1841 CSD10
                          |            |         M. lineatipes R55
                          |            |         M. rangii B88
                          |            |         M. rotundata DH02
                          |            |--M. (Chalicodomopsis) Engel 1999 E01
                          |            |    `--M. (*C.) glaesaria Engel 1999 E01
                          |            `--M. (Mitchellapis) fabricator YHT05
                          `--+--Chelostoma Latreille 1809 CSD10, E01
                             |    |--C. californicum Cresson 1878 CSD10
                             |    `--C. mauritanica Lucas 1849 E12
                             `--+--+--Protosmia rubifloris (Cockerell 1898) CSD10
                                |  `--Heriades Spinola 1808 CSD10, E01 (see below for synonymy)
                                |       `--H. crucifer Cockerell 1897 CSD10
                                `--+--Hoplitis albifrons (Kirby 1837) CSD10
                                   `--+--Ashmeadiella aridula Cockerell 1910 CSD10
                                      `--+--Hoplosmia scutellaris (Morawitz 1868) CSD10
                                         `--Osmiina E01
                                              |--Metalinella E01
                                              `--Osmia Panzer 1806 CSD10, E01
                                                   |--O. cornifrons KIW98
                                                   |--O. lignaria Say 1837 CSD10
                                                   |--O. luctuosa Lucas 1848 E12
                                                   |--O. metallica Lucas 1848 E12
                                                   `--O. pumila KIW98

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

Glyptapis fuscula Cockerell 1909 [incl. G. neglecta Salt 1931, G. reducta Cockerell 1909, G. reticulata Cockerell 1909] E01

Heriades Spinola 1808 CSD10, E01 [incl. Trypetes Schenck 1861 E01; Heriadini, Trypetina]

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[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.

[DH02] Donovan, B. J., A. Havron, D. M. Leathwick & J. S. Ishay. 2002. Release of Sphecophaga orientalis Donovan (Hymenoptera: Ichneumonidae: Cryptinae) in New Zealand as a possible ‘new association’ biocontrol agent for the adventive social wasps Vespula germanica (F.) and Vespula vulgaris (L.) (Hymenoptera: Vespidae: Vespinae). New Zealand Entomologist 25: 17-25.

[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.

[KIW98] Kearns, C. A., D. W. Inouye & N. M. Waser. 1998. Endangered mutualisms: the conservation of plant-pollinator interactions. Annual Review of Ecology and Systematics 29: 83-112.

[OC01] OConnor, B. M. 2001. Historical ecology of the Acaridae (Acari): Phylogenetic evidence for host and habitat shifts. In Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) pp. 76-82. CSIRO Publishing: Melbourne.

[R55] Rayment, T. 1955. Taxonomy, morphology and biology of sericophorine wasps with diagnoses of two new genera and descriptions of forty new species and six sub-species. Memoirs of the National Museum, Melbourne 19: 1-95.

[YHT05] Yates, C. J., S. D. Hopper & R. H. Taplin. 2005. Native insect flower visitor diversity and feral honeybees on jarrah (Eucalyptus marginata) in Kings Park, an urban bushland remnant. Journal of the Royal Society of Western Australia 88 (4): 147-153.

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. Despite their small size, a number of species 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).

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 [incl. Schizopelta McAreavey 1949 H09]
    |--*M. (Monomorium) minutum Mayr 1855 [incl. Myrmica (Monomorium) carbonaria Smith 1858] B03
    `--M. (Chelaner Emery 1914) E14
         |--M. (C.) forcipatum Emery 1914 E14
         `--M. (C.) longipes Emery 1914 E14

Monomorium incertae sedis:
  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. atomus Forel 1902 [incl. M. atomus var. integrius] B03
  M. australicum H09 [incl. M. talpa FT08]
  M. barbatulum E89
  M. bicorne [incl. M. macareaveyi] H09
  M. bihamatum H09
  M. brachythrix H09
  M. centrale H09
  M. crinitum H09
  M. croceiventre Emery 1914 E14
  M. decuria H09
  M. destructor (Jerdon 1851) (see below for synonymy) B03
  M. dichroum Forel 1902 B03
  M. disetigerum H09
  M. durokoppinense H09
  M. edentatum Emery 1897 E14
  M. elegantulum H09
  M. eremophilum H09
  M. ergatogyna BS06
  M. euryodon H09
  M. falcatum [=*Schizopelta falcata] H09
  M. fieldi [incl. M. laeve nigrium] H09
  M. flavonigrum H09
  M. floricola (Jerdon 1851) [=Atta floricola; incl. M. specularis Mayr 1866] B03
  M. fossulatum Emery 1894 B03
  M. glyciphilum (Smith 1858) [=Myrmica glyciphila] B03
  M. gracillimum (Smith 1861) [=Myrmica gracillima; incl. Mo. gracillimum var. mayri Forel 1902] B03
  M. hildebrandti H09
  M. indicum Forel 1902 [=M. salomonis indicum] B03
  M. integrum E14
  M. kilianii H09
  M. lacunosum H09
  M. laeve H09
  M. latinode Mayr 1872 FT08
  M. leae [incl. M. hemiphaeum] H09
  M. legulus H09
  M. longi Forel 1902 B03
  M. longiceps H09
  M. longinode H09
  M. luteum G89
  M. majeri H09
  M. megalops H09
  M. melleum Emery 1914 E14
  M. micula H09
  M. monomorium H09
  M. nanum H09
  M. nitidum E14
  M. orientale Mayr 1879 FT08
  M. pergandei [=Epoecus pergandei] WW90
  M. pharaonis (Linnaeus 1758) (see below for synonymy) B03
  M. pubescens H09
  M. rothsteini [=Chelaner rothsteini] H09
  M. rubriceps E14
  M. rufiventris [=Vollenhovia rufiventris] E14
  M. rufonigrum H09
  M. sagei Forel 1902 B03
  M. sculpturatum H09
  M. silaceum H09
  M. sordidum [=Chelaner sordidum] H09
  M. stictonotum H09
  M. striatifrons H09
  M. subcoecum Emery 1894 FT08
  M. sublamellatum H09
  M. sydneyense H09
  M. tricolor Emery 1914 E14
  M. turneri [=Vollenhovia turneri] E14
  M. wroughtoni Forel 1902 B03
  M. xantheklemma H09

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

Monomorium pharaonis (Linnaeus 1758) [=Formica pharaonis; incl. Myrmica domestica Schuck. 1838, Atta minuta Jerdon 1851] B03

*Type species of generic name indicated

REFERENCES

[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera. Vol. II. 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 2a 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, Heft IV 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 2a 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.

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

[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.

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 Europe and the Americas (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 [Nothomyrmecii] WB03a
    |    |--Prionomyrmex Mayr 1868 WB03a
    |    |    |--P. janzeni WB03a
    |    |    `--P. longiceps WB03a
    |    `--+--Nothomyrmecia Clark 1934 WB03a
    |       |    `--N. macrops WB03a
    |       `--Archimyrmex WB03b (see below for synonymy)
    |            |--*A. rostratus WB03b
    |            |--‘Ameghinoia’ piatnitzkyi WB03b
    |            `--‘Polanskiella’ smekali WB03b
    `--Myrmecia Fabricius 1804 (see below for synonymy) WB03a
         |--M. acuta H09
         |--M. analis H09
         |--M. apicalis Emery 1883 E14
         |--M. arnoldi H09
         |--M. callima H09
         |--M. cephalotes H09
         |--M. chasei H09
         |--M. clarki H09
         |--M. croslandi MG06
         |--M. desertorum H09
         |--M. dispar H09
         |--M. elegans H09
         |--M. erecta H09
         |--M. forceps H09
         |--M. forficata B88
         |--M. fulgida H09
         |--M. fulvipes WB03a
         |--M. fuscipes H09
         |--M. gratiosa H09
         |--M. gulosa H09
         |--M. hilli H09
         |--M. infima H09
         |--M. inquilina H09
         |--M. ludlowi H09
         |--M. mandibularis H09
         |--M. michaelseni H09
         |--M. mjobergi WB03a
         |--M. nigra H09
         |--M. nigriceps H09
         |--M. nigriscapa H09
         |--M. nigrocincta ZS10
         |--M. nobilis WB03a
         |--M. occidentalis H09
         |--M. pavida H09
         |--M. picta WB03a
         |--M. picticeps H09
         |--M. piliventris MS04
         |--M. pilosula WB03a
         |--M. pyriformis WB03a
         |--M. regularis WB03a
         |--M. rubripes H09
         |--M. rugosa H09
         |--M. swalei H09
         |--M. tarsata WB03a
         |--M. tepperi H09
         |--M. testaceipes H09
         |--M. urens H09
         |--M. varians WB03a
         `--M. vindex H09

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

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[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, Heft IV, 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.

[MS04] Majer, J. D., S. O. Shattuck, A. N. Andersen & A. J. Beattie. 2004. Australian ant research: fabulous fauna, functional groups, pharmaceuticals, and the Fatherhood. Australian Journal of Entomology 43 (3): 235–247.

[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.

[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.

[ZS10] Zborowski, P. & R. Storey. 2010. A Field Guide to Insects in Australia. Reed New Holland: Sydney.

Dolichoderinae

Long-necked ant Dolichoderus attelaboides, photographed by Tom Murray.


Belongs within: Formicidae.
Contains: Iridomyrmex, Technomyrmex, Tapinoma.

The Dolichoderinae are a group of ants characterised by the presence of a transverse slit at the apex of the abdomen from which chemicals are secreted (Heterick 2009). A number of dolichoderines are notably aggressive, and some such as the Argentine ant Linepithema humile and white-footed ant Technomyrmex albipes are notorious pests.

Characters (from Heterick 2009): Mandibles triangular with teeth along entire inner margin; abdomen without impression between third and fourth abdominal segments, often soft, flexible and easily collapsed; transverse slit present at junction of hypopygium and pygidium.

<==Dolichoderinae [Dolichoderidae]
    |--Dolichoderus Lund 1831 BS06, B03
    |    |--*D. attelaboides B03
    |    |--D. affinis Emery 1889 [incl. D. affinis var. glabripes, D. affinis var. nigricans] E89b
    |    |--D. angusticornis H09
    |    |--D. australis E14
    |    |--D. bituberculatus (Mayr 1862) B03 [=Hypoclinea bituberculata B03; incl. Tapinoma gibba E89b]
    |    |--D. burmanicus Bingham 1903 B03
    |    |--D. clusor H09
    |    |--D. cuspidatus N-ZLI11
    |    |--D. feae Emery 1889 E89b
    |    |--D. formosus H09
    |    |--D. fuscus Emery 1889 [=D. feae fuscus] B03
    |    |--D. gibbifer E89b
    |    |--D. glauerti H09
    |    |--D. gracilipes [incl. D. semirufus André 1887] E89b
    |    |--D. moggridgei Forel 1880 B03
    |    |--D. nigricornis H09
    |    |--D. occidentalis H09
    |    |--D. quadripunctatus [incl. D. sibiricus] E89a
    |    |--D. reflexus H09
    |    |--D. scabridus BS06
    |    |--D. sulcaticeps (Mayr 1870) [=Hypoclinea sulcaticeps] B03
    |    |--D. taprobanae (Smith 1858) (see below for synonymy) B03
    |    |--D. thoracicus N-ZLI11 [=Tapinoma thoracica E89b]
    |    |--D. tricolor Emery 1914 E14
    |    |--D. tuberifer E89b
    |    `--D. ypsilon H09
    |         |--D. y. ypsilon H09
    |         |--D. y. nigra H09
    |         `--D. y. rufotibialis H09
    `--+--+--Technomyrmex BS06
       |  `--+--Tapinoma BS06
       |     `--Liometopum Mayr 1861 BS06, B03
       |          |--*L. microcephalum B03
       |          |--L. apiculatum BS06
       |          |--L. lindgreeni Forel 1902 B03
       |          `--L. occidentale BS06
       |               |--L. o. occidentale BM76
       |               `--L. o. luctuosum BM76
       `--+--Leptomyrmex BS06
          |    |--L. erythrocephalus BS06
          |    `--L. pallens Emery 1883 [incl. L. p. var. geniculata Emery 1914, L. p. var. nigriceps Emery 1914] E14
          `--+--+--Forelius pruinosus BS06 [=Iridomyrmex pruinosus WW90]
             |  |    |--F. p. pruinosus BM76
             |  |    `--‘Iridomyrmex’ p. analis BM76
             |  `--Dorymyrmex BS06
             |       |--D. bicolor BS06 [=Conomyrma bicolor WW90, D. pyramicus bicolor WW90]
             |       `--D. pyramicus BM76
             `--+--Azteca BS06
                |    |--A. ovaticeps BS06
                |    `--A. xanthochroa WI-B09
                `--+--Linepithema humile BS06
                   `--+--Papyrius nitidus BS06
                      `--+--+--Philidris cordatus BS06
                         |  `--Turneria bidentata BS06
                         `--Anonychomyrma BS06
                              |--A. gilberti BS06
                              |--A. itinerans H09
                              |    |--A. i. itinerans H09
                              |    `--A. i. perthensis H09
                              `--A. nitidiceps H09

Dolichoderinae incertae sedis:
  Iridomyrmex H09
  Eotapinoma GE05
    |--E. macalpini GE05
    `--E. zherichinius GE05
  Ochetellus Shattuck 1992 FT08
    |--O. flavipes H09
    |--O. glaber (Mayr 1862) FT08 [=Hypoclinea glabra B03, Iridomyrmex glaber FT08]
    `--O. punctatissimus H09
  Froggattella H09
    |--F. kirbii H09
    `--F. latispina H09
  Arnoldius H09
    |--A. flavus H09
    `--A. scissor H09
  Nebothriomyrmex majeri H09
  Doleromyrma H09
    |--D. darwiniana WFS04
    |    |--D. d. darwiniana H09
    |    `--D. d. fida H09
    `--D. rottnestensis [=Tapinoma minutum rottnestense] H09
  Hypoclinea cuspidatus H09, GE05
  Conomyrma WW90
    |--C. flavopectus [=Dorymyrmex pyramicus flavopectus] WW90
    `--C. insana WW90
  Bothriomyrmex Emery 1865 B03
    |--*B. meridionalis B03
    |--B. dalyi Forel 1895 [=B. wroughtoni dalyi] B03
    |--B. decapitans A71
    |--B. myops Forel 1895 B03
    |--B. walshi Forel 1895 B03
    `--B. wroughtoni Forel 1895 B03

Dolichoderus taprobanae (Smith 1858) [=Formica taprobanae; incl. Hypoclinea gracilipes Mayr 1878, Dolichoderus taprobanae var. gracilipes, D. semirufus André 1887] B03

*Type species of generic name indicated

REFERENCES

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

Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera. Vol. II. Ants and Cuckoo-Wasps. Taylor and Francis: London.

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

[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.

[E89a] Emery, C. 1889a. Intorno ad alcune formiche della fauna palearctica. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 439–443.

[E89b] Emery, C. 1889b. 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.

[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, Heft IV, 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.

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

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

[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.

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.

Wagner, R., & S. Ibáñez-Bernal. 2009. Psychodidae (sand flies, and moth flis or owl 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. 319–335. Ottawa, NRC Research Press.

[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.

[WFS04] Winks, C. J., S. V. Fowler & L. A. Smith. 2004. Invertebrate fauna of boneseed, Chrysanthemoides monilifera ssp. monilifera (L.) T. Norl. (Asteraceae: Calenduleae), an invasive weed in New Zealand. New Zealand Entomologist 27: 61–72.