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Subfamily MYRMICINAE
Tribe CREMATOGASTRINI

Tetramorium bicarinatum (Nylander, 1846)

Bicolored pavement ant

Uploaded 2009; last updated 29 March 2016
Authors: Joe A. MacGown and Ryan J. Whitehouse

Tetramorium bicarinatum, full face view of worker (MS, Jackson Co.) (photo by Joe A. MacGown)
Tetramorium bicarinatum, lateral view of worker (MS, Jackson Co.) (photo by Joe A. MacGown)
Tetramorium bicarinatum, lateral view of worker (MS, Jackson Co.) (photo by Joe A. MacGown)
Tetramorium bicarinatum, full face view of worker (MS, Pearl River Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, lateral view of worker (MS, Pearl River Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, dorsal view of worker (MS, Pearl River Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, full face view of dealate queen (CA, Los Angeles Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, lateral view of dealate queen (CA, Los Angeles Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, dorsal view of dealate queen (CA, Los Angeles Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, full face view of alate queen (MS, Pearl River Co.) (photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, lateral view of alate queen (MS, Pearl River Co.)
(photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, dorsal view of alate queen (MS, Pearl River Co.)
(photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, dorsal view of alate queen (MS, Pearl River Co.)
(photo by Ryan J. Whitehous and Joe A. MacGown)
Tetramorium bicarinatum, full face view of male (CA, Los Angeles Co.) (photo by AntWeb.org)
Tetramorium bicarinatum, lateral view ofmale (CA, Los Angeles Co.) (photo by AntWeb.org)
Tetramorium bicarinatum, dorsal view of male(CA, Los Angeles Co.) (photo by AntWeb.org)

Introduction
Tetramorium species can be found worldwide with the largest diversity being in Africa. Human commerce and travel have greatly aided in the movement of some members of this genus around the world where several species have successfully established non-native populations. In their non-native ranges, Tetramorium species are often found in urban environments, including insides structures, and may be the most common species present.

Tetramorium species can be identified by the two segmented waist; broad frons; 11 or 12-segmented antennae; three-segmented antennal club; antennal scrobe present dorsal to the eyes; eyes located along the midline of the head; lateral part of the clypeus forming a sharp wall anterior to the antennal insertion; and the sting often with a lamellate appendage found apicodorsally that projects at an angle to the long axis of the sting shaft.

Tetramorium bicarinatum is distrubuted worldwide and is one of the most abundant species of Tetramorium. It is known to occure in tropical and subtropical climates and has been found in heated buildings in more temperate areas. Historically this species was misidentified as T. guineese (Fabricius, 1793).

Taxonomic History
Myrmica bicarinata Nylander, 1846: 1061 (w.q.) U.S.A. Nearctic. Forel, 1891: 151 (m., misidentified as Tetramorium guineense); Wheeler & Wheeler, 1954: 449 (l., misidentified as Tetramorium guineense); Wheeler, 1924: 136 (gynandromorph, misidentified as Tetramorium guineense). Combination in Tetramorium: Mayr, 1862: 740; Bolton, 1977: 94. Junior synonym of Formica guineensis Fabricius, 1793: 357 (now in Pheidole): Mayr, 1862: 740. Revived from synonymy and senior synonym of Tetramorium cariniceps (and its junior synonym Tetramorium kollari), Tetramorium modesta Smith, Tetramorium reticulata: Bolton, 1977: 94. [Note. The names Tetramorium cariniceps, Tetramorium kollari and Tetramorium reticulata had previously been incorrectly synonymised with Tetramorium guineensis Fabricius by Roger, 1862: 293; Tetramorium modesta Smith was wrongly synonymised with Tetramorium guineensis by Donisthorpe, 1932: 463.]. See also: Bolton, 1979: 164; Hita Garcia & Fisher, 2011: 18.

Identification
Worker: HL 0.90-0.97mm, HW 0.77-0.81mm, SL 0.60-0.64mm, EL 0.21-0.23mm, MeSL 0.97-1.01mm (n=5) (MEM specimens). Head, mesosoma, and waist orange to orangish brown; legs lighter, yellowish brown; and gaster dark brown. Head strongly shining, with strong, medial, longitudinal rugae extending from clypeus towards the posterior border, strong rugoreticulation present on posterior region of head (in full face view) and on sides of head; numerous erect setae present; eyes distinctly convex and situated along the midline of the head; mandibles triangular in shape with six teeth along the inner margin; anterior clypeal margin distinctly notched or indented; lateral portions of clypeus forming a sharp wall anterior to antennal insertion point; antennae 12-segmented with a three-segmented club; scape fits against head along a longitudinal ridge running from the posterior end of the clypeus to the posterior border of the head. Mesosoma shiny, with strong rugoreticulation, especially dorsally, becoming transverse rugue on mesopleuron and metapleuron; erect setae present on dorsal surface; dorsal outline relatively flat and continuous in profile; propodeal spines well developed and bidentate with the length of each dorsal spine about twice the length of each ventral one. Waist two-segmented with prominent nodes, both nodes with strong reticulation; with conspicuous, erect setae; petiolar node and postpetiolar node subequal in length; postpetiolar node wider than petiolar node; petiole with a short tooth present anteroventrally, tooth directed slightly anteriorly. Gaster shining with conspicuous, erect setae; first tergite makes up at least half of length; sting with a lamellate appendage found apicodorsally that projects at an angle to the long axis of the sting shaft.

Queen: HL 1.07-1.12mm, HW 0.92-0.95mm, SL 0.66-0.69mm, EL 0.28-0.30mm, MeSL 1.45-1.51mm (n=5) (MEM specimens). Head, mesosoma, and waist orange reddish brown; legs light, orangish brown; and gaster dark brown. Head strongly shining, with medial, longitudinal rugae extending from clypeus towards the posterior border, strong rugoreticulation present on posterior region of head (in full face view) and on sides of head; numerous erect setae present; eyes well developed and situated along the midline of the head; three whitish colored ocelli present; mandibles triangular; clypeus notched or indented along the anterior border; lateral portions of clypeus forming a sharp wall anterior to antennal insertion point; antennae 12-segmented with a three-segmented club; scape fits against head along a longitudinal ridge running from the posterior side of the clypeus to the posterior border of the head. Mesosoma shiny; pronotum and propodeum with strong rugoreticulation; mesoscutellum and mesoscutum with longitudinal rugue; anepisternum with transverse rugue, katepisternum moslty lacking sculpture except at edges; numerous simple, erect setae present on mesosomal dorsum; dorsal surface flat with a distinct notch at the propodeal suture; propodeal spines bidentate with the length of each dorsal spine about twice the length of the each ventral spine; mesosoma enlarged with four wings or wing scars; wings, when present, hylaline, with typical myrmicine venation; forewing with pale stigma, elongate costal cell, basal cell, one submarginal, marginal cell, one discal cell, and subbasal cell; medial vein almost reaches wing tip; hindwing hindwing lacking jugal lobe and with costal, basal, and subbasal cells closed. Waist two-segmented, both nodes with strong reticulation; conspicuous, posteriorly directed, erect setae present; petiolar node and postpetiolar node subequal in length: petiole with a short tooth present anteroventrally, tooth directed slightly anteriorly. Gaster shining with conspicuous, erect setae; first tergite makes up at least half of length; sting present and with a lamellate appendage found apicodorsally that projects at an angle to the long axis of the sting shaft.

Male: (no MEM specimens, based on antweb.org photos). Head fading from yellowish brown anteriorly to brown posteriorly, mesosoma, waist, and gaster grown with lighter yellowish brown coloration at posterior edges of gastral segments; antennae and legs pale yellow brown. Head with longitudinal rugae medially extending from anterior portion of clypeus to base of lower ocellus, remainder of head with strong, rugureticulation, with fine punctulation present in spaces formed between; numerous erect setae present of various lengths, with most of the setae on posterior border being longer than those on the face; eyes large, about half the length of the head; three large, whitish colored ocelli present, slightly raised; mandibles triangular with elongate apical tooth followed by four smaller teeth; anteriorly clypeal border relatively straight; antennae ten-segmented; scape and second funicular segment subequal in length. Mesosoma not overly shiny, slightly matte in appearance, mostly lacking sculpture, except propodeum, which has loosely connected rugureticulum, and transverse sculpture present on scutellar sulcus and other deep sutural lines; scattered semi erect setae present, especially dorsally; mesoscutem enlarged, smoothly curved and overhanging propodeum; wings transparent; with similar venation to female, except pterostigman in forewing mostly transparent. Propodeal spines lacking. Waist two-segmented with scattered, posteriorly directed, erect setae; petiole with loose rugoreticulation present; a sharp subpetiolar process directed anteriorly present anterioventrally; petiole elongate, longer than postpetiole; postpetiole somewhat circular. Gaster lacking obvious sculpture, with conspicuous, erect setae; first tergite darker than the remaining ones; genitalia visible at the apex, in side view parameres triangular, rounded apically.

Tatramorium bicarinatum can be separated from other Tetramorium species in the Southeastern United states by its relatively flat mesosoma, the medial indentation of the clypeus; its bicoloration; and strong rugoreticulation on head, mesosoma, and waist.

Biology and Economic Importance
Tetramorium bicarinatum may bethe most abundant ant in its genus based on its worldwide distribution in tropical regions. However, it does not have a strong presence in the Afrotropical region, possibly as a result of competition from other Tetramorim species. Tetramorium bicarinatum is a common ant along the Gulf Coast and has been frequently found on plants in quarantine (Smith 1965). This ant can also be found farther north in heated buildings, especially greenhouses, where the artificially stable temperatures are closer to their preferred temperature range. Tetramorium bicarinatum has been collected on palmettos and plant nurseries in Mississippi and has been reported to form colonies under rocks, tree bark, exposed soil, rotting logs and other objects found on the ground. They are also known to like honeydew producing insects and reported to feed on dead and live insects (Smith 1965).

Although Tetramorium bicarinatum can occasionally be found nesting in building and houses, it is not considered to be an important pest species. When in homes they are most probably omnivorous and will feed on grease, meats, sugary food and plant based foods (Smith 1965).

Distribution
Native Range: Southeast Asia (Bolton 1977, 1979, 1980)

Afrotropical Region: Cape Verde, Comoros, Eritrea, Ghana, Macaronesia, United Arab Emirates, Yemen (antweb.org and antwiki.org).
Australasian Region: Australia, Lord Howe Island, New Caledonia, New Guinea, New Zealand, Norfolk Island, Solomon Islands (antweb.org and antwiki.org).
Indo-Australian Region: American Samoa, Borneo, Fiji, French Polynesia, Guam, Hawaii, Indonesia, Kiribati, Krakatau Islands, Malaysia, Marshall Islands, Micronesia (Federated States of), Niue, Philippines, Samoa, Tokelau, Tonga, Vanuatu (antweb.org and antwiki.org).
Malagasy Region: Europa Island, Madagascar, Mauritius, Mayotte, Reunion, Seychelles (antwiki.org).
Nearctic Region: United States (type locality) (antwiki.org).
Neotropical Region: Aruba, Barbados, Brazil, British Virgin Islands, Costa Rica, Cuba, Dominican Republic, Ecuador, French Guiana, Galapagos Islands, Greater Antilles, Guyana, Haiti, Honduras, Martinique, Mexico, Netherlands Antilles, Nicaragua, Panama, Peru, Puerto Rico, Trinidad and Tobago (antweb.org and antwiki.org).
Oriental Region: Bangladesh, Bhutan, Borneo, India, Nicobar Island, Sri Lanka, Thailand, Vietnam (antweb.org and antwiki.org).
Palearctic Region: Austria, Belgium, Canary Islands, China, Denmark, Germany, Greece, Hungary, Iberian Peninsula, Israel, Japan, Montenegro, Netherlands, Republic of Korea, Spain, Sweden (antweb.org and antwiki.org).

U.S. Distribution: AL, AR, CA, FL, GA, HI, IL, LA, MS, MO, NC, SC, TX (antweb.org and MEM).
Southeastern U.S. Distribution: AL, FL, GA, LA, MS, NC, SC (MEM).

Acknowledgments
Thanks to Ryan J. Whitehouse for help with measuring specimens, comments on descriptions, and proofreading. Funding for the ant work being done by the MEM in Alabama and Mississippi is from several sources including the National Institute of Food and Agriculture, United States Department of Agriculture, under Project No. MIS-012040, the Mississippi Agricultural and Forestry Experiment Station at Mississippi State University, with support from State Project MIS-311080, NSF Grants BSR-9024810 and DFB-9200856, the Tombigbee National Forest (U.S. Forest Service), the Noxubee Wildlife Refuge, Mississippi Natural Heritage Program Research Grant, USDA Forest Service Agreement No. 08-99-07-CCS-010, the William H. Cross Expedition Fund, and primarily by the USDA-ARS Areawide Management of Imported Fire Ant Project. Additionally, special cooperation has been provided by State Parks, National Forests, National Wildlife Refuges, the Natchez Trace Parkway, and from various private landowners in both Alabama and Mississippi.

Literature Cited
Bolton, B. 1977. The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Oriental and Indo-Australian regions, and in Australia. Bulletin of the British Museum (Natural History). Entomology 36:67-151.

Bolton, B. 1979. The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Malagasy region and in the New World. Bulletin of the British Museum (Natural History). Entomology 38:129-181.

Bolton, B. 1980. The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Ethiopian zoogeographical region. Bull. Br. Mus. (Nat. Hist.) Entomol. 40: 193-384

Donisthorpe, H. 1932. On the identity of Smith's types of Formicidae (Hymenoptera) collected by Alfred Russell Wallace in the Malay Archipelago, with descriptions of two new species. Annals and Magazine of Natural History (10)10:441-476.

Fabricius, J. C. 1793. Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species, adjectis synonimis, locis observationibus, descriptionibus. Tome 2. Hafniae [= Copenhagen]: C. G. Proft, 519 pp.

Forel, A. 1891. Les Formicides. [part]. In: Grandidier, A. 1891. Histoire physique, naturelle, et politique de Madagascar. Volume XX. Histoire naturelle des Hyménoptères. Deuxième partie (28e fascicule). Paris: Hachette et Cie, v + 237 pp.

Hita Garcia, F.; Fisher, B. L. 2011. The ant genus Tetramorium Mayr (Hymenoptera: Formicidae) in the Malagasy region—introduction, definition of species groups, and revision of the T. bicarinatumT. obesumT. sericeiventre and T. tosii species groups. Zootaxa 3039:1-72.

Mayr, G. 1862. Myrmecologische Studien. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 12:649-776.

Nylander, W. 1846. Additamentum adnotationum in monographiam formicarum borealium Europae. Acta Societatis Scientiarum Fennicae 2:1041-1062.

Roger, J. 1862. Synonymische Bemerkungen. 1. Ueber Formiciden. Berliner Entomologische Zeitschrift 6:283-297

Smith, M.R. 1965. House-infesting ants of the Eastern United States, Their Recognition, Biology, and Economic Importance. Agricultural Research Service, United States Department of Agriculture Technical Bulletin No. 1326.

Wheeler, W. M. 1924. A gynandromorph of Tetramorium guineense Fabr. Psyche (Cambridge) 31:136-137.

Wheeler, G. C.; Wheeler, J. 1954. The ant larvae of the myrmicine tribes Meranoplini, Ochetomyrmicini and Tetramoriini. American Midland Naturalist 52:443-452.

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