Temnothorax nevadensis

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Temnothorax nevadensis
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmicinae
Tribe: Crematogastrini
Genus: Temnothorax
Species group: andrei
Species: T. nevadensis
Binomial name
Temnothorax nevadensis
(Wheeler, W.M., 1903)

Temnothorax-nevadensis-MCZ001L.jpg

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Specimen Label

Synonyms

Modified from Mackay (2000): This species nests in soil in moist areas, generally under stones or in rotten wood. It may be involved in plesiobiosis, as Wheeler (l903a) reported it nesting at the entrance of a Mycetomoellerius turrifex nest and at the edge of a flourishing colony of Pheidole tepicana (= P. instabilis).

Identification

Prebus 2017 - A member of the andrei clade.

Mackay (2000) - This species is fairly common throughout the western United States. The clypeus has a moderately well defined medial carina and 2 well-defined lateral carina, which curve and cross the anterior part of the medial lobe of the clypeus. It has a completely and strongly striated head with punctures between the striae, and with rarely a slender median strip without sculpture. The sides of the pronotum are distinctly and coarsely rugose, with the background weakly punctate, but shining. The dorsum of the mesonotum is striate to finely rugose, but mixed with background punctures and not as shiny as the background of the sides of the pronotum. The petiole and postpetiole are primarily punctate, but fine rugulae can be seen on the dorsum of the petiole. There is usually a single ruga on the side of the petiole. Some larger specimens have several poorly defined rugae on the side of the petiole (making it look "wrinkled"), and occasionally on the anterior face. These larger specimens usually have longer propodeal spines, sometimes as long as the distance between the bases. These specimens were referred to as Temnothorax nevadensis subsp. melanderi in the past.

Temnothorax nevadensis and Temnothorax neomexicanus have similar lateral clypeal carinae, which usually curve and connect on the anterior part of the medial lobe of the clypeus. They are thus apparently closely related. They can be easily separated as the dorsum of the head of Temnothorax nevadensis is nearly completely sculptured, whereas part of the head of Temnothorax neomexicanus is smooth and shining.

Keys including this Species

Distribution

Western US including Washington, Idaho, Montana, Wyoming, California, Nevada, Colorado, Arizona (Navajo Reservation) and Utah. Mexico: Baja California and Baja California Sur. Canada: SW British Columbia.

Latitudinal Distribution Pattern

Latitudinal Range: 50.65° to 30.98555556°.

   
North
Temperate
North
Subtropical
Tropical South
Subtropical
South
Temperate

Distribution based on Regional Taxon Lists

Nearctic Region: United States (type locality).
Neotropical Region: Mexico.

Distribution based on AntMaps

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Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
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Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.
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Habitat

Ward (2005) "occupying a broad range of habitats, including coastal scrub, chaparral, oak woodland, open coniferous forest, and sagebrush desert."

Wheeler and Wheeler (1986) "Known from cool desert, pinyon-juniper forest, coniferous forest and alpine areas."

Mackay (2000) "D. S. Chandler (pers. comm.) collected this species in litter in a number of plant communities, including tanbark oak, oak leaf litter near a spring, maple and oak litter, Douglas fir, and laurel.

Mackay and Mackay (2002) - Nests in soil in moist areas, usually under stones, or in rotten wood, from communities including cool desert, pinyon- juniper, coniferous forest and alpine areas. Specimens were collected in litter in a number of plant communities, including tanbark oak, oak leaf litter near a spring, maple and oak litter, Douglas fir, and laurel.

Abundance

Commmon.

Biology

Castes

Worker

Mcz-ent00669292 Temnothorax nevadensis hef.jpgMcz-ent00669292 Temnothorax nevadensis hal.jpgMcz-ent00669292 Temnothorax nevadensis had.jpgMcz-ent00669292 Temnothorax nevadensis lbs.JPG
. Owned by Museum of Comparative Zoology.

Images from AntWeb

Temnothorax nevadensis casent0103165 head 1.jpgTemnothorax nevadensis casent0103165 profile 1.jpgTemnothorax nevadensis casent0103165 dorsal 1.jpgTemnothorax nevadensis casent0103165 label 1.jpg
Paratype of Temnothorax nevadensisWorker. Specimen code casent0103165. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by USNM, Washington, DC, USA.
Temnothorax nevadensis casent0103385 head 1.jpgTemnothorax nevadensis casent0103385 profile 1.jpgTemnothorax nevadensis casent0103385 dorsal 1.jpgTemnothorax nevadensis casent0103385 label 1.jpg
Paratype of Temnothorax nevadensisWorker. Specimen code casent0103385. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by USNM, Washington, DC, USA.
Temnothorax nevadensis casent0103389 head 1.jpgTemnothorax nevadensis casent0103389 profile 1.jpgTemnothorax nevadensis casent0103389 dorsal 1.jpgTemnothorax nevadensis casent0103389 label 1.jpg
Paratype of Temnothorax nevadensisWorker. Specimen code casent0103389. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by USNM, Washington, DC, USA.
Temnothorax nevadensis casent0103390 head 1.jpgTemnothorax nevadensis casent0103390 profile 1.jpgTemnothorax nevadensis casent0103390 dorsal 1.jpgTemnothorax nevadensis casent0103390 label 1.jpg
Paratype of Temnothorax nevadensisWorker. Specimen code casent0103390. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by USNM, Washington, DC, USA.
Temnothorax nevadensis casent0005685 head 1.jpgTemnothorax nevadensis casent0005685 profile 1.jpgTemnothorax nevadensis casent0005685 dorsal 1.jpgTemnothorax nevadensis casent0005685 label 1.jpg
Worker. Specimen code casent0005685. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by UCDC, Davis, CA, USA.

Queen

Images from AntWeb

Temnothorax nevadensis casent0103166 head 1.jpgTemnothorax nevadensis casent0103166 profile 1.jpgTemnothorax nevadensis casent0103166 dorsal 1.jpgTemnothorax nevadensis casent0103166 label 1.jpg
Paratype of Temnothorax nevadensisQueen (alate/dealate). Specimen code casent0103166. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by USNM, Washington, DC, USA.
Mcz-ent00668939 Temnothorax nevadensis queen hef.jpgMcz-ent00668939 Temnothorax nevadensis queen hal.jpgMcz-ent00668939 Temnothorax nevadensis queen had.jpgMcz-ent00668939 Temnothorax nevadensis queen lbs.JPG
. Owned by Museum of Comparative Zoology.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • nevadensis. Leptothorax nevadensis Wheeler, W.M. 1903c: 252, pl. 12, fig. 20 (w.q.m.) U.S.A. Wheeler, G.C. & Wheeler, J. 1973b: 71 (l.). Combination in L. (Myrafant): Smith, D.R. 1979: 1393; in Temnothorax: Bolton, 2003: 271. Senior synonym of eldoradensis, melanderi: Mackay, 2000: 376; of lindae, maryanae: Ward, 2005: 16.
  • melanderi. Leptothorax melanderi Wheeler, W.M. 1909e: 81 (w.) U.S.A. Subspecies of nevadensis: Creighton, 1950a: 266. Junior synonym of nevadensis: Mackay, 2000: 376.
  • eldoradensis. Leptothorax eldoradensis Wheeler, W.M. 1915b: 414 (w.) U.S.A. Subspecies of nevadensis: Creighton, 1950a: 266. Junior synonym of nevadensis: Mackay, 2000: 376.
  • lindae. Leptothorax (Myrafant) lindae Mackay, 2000: 356, figs. 51, 52, 123, 124 (w.q.) U.S.A. Combination in Temnothorax: Bolton, 2003: 271. Junior synonym of nevadensis: Ward, 2005: 16.
  • maryanae. Leptothorax (Myrafant) maryanae Mackay, 2000: 364, figs. 34, 42, 129-131 (w.q.m.) U.S.A. Combination in Temnothorax: Bolton, 2003: 271. Junior synonym of nevadensis: Ward, 2005: 17.

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Description

Worker

Length 2.5-3 mm. Mandibles 5-toothed. Clypeus depressed in the middle, its anterior margin sinuately excised. Antennae 12-jointed, scape reaching to posterior corner of the head, first funicular joint as long as joints 2-4 together, joints 2-6 slightly broader than long, joints 7 and 8 as long as broad; two basal joints of club subequal, together shorter than the terminal joint. Thorax above in front of about the same width as below and behind; humeri much rounded, dorsum flattened in profile,without mesoepinotal constriction. Epinotal spines robust, pointed, decidedly longer than broad at their bases, and nearer together at their bases than long, directed upward, outward and backward. Petiole 1 1/2 times as long as broad, sides of node rounded, broader than the peduncle; .seen from above the node is transverse, in profile it is narrow antero-posteriorly, its anterior slope gently ascending, concave, its posterior slope more abrupt, also concave, summit rounded; ventral surface of petiole with a prominent, compressed, downwardly directed tooth. Postpetiole in profile with a prominent, sometimes slightly angular node; the segment seen from above is transversely elliptical, about half again as broad as long, its anterior angles rounded. Gaster and legs of the usual conformation.

Mandibles striate and punctate. Clypeus subopaque, its surface, especially at the sides, traversed by rather coarse longitudinal rugaae. Head with a satiny luster, sparsely punctate and with delicate longitudinal rugalae which become decidedly reticulate in the antennal foveae. Thorax subopaque, its dorsal surface irregularly reticulate-rugose and foveolate, pleurae, petiole and postpetiole regularly foveolate-punctate; posterior epinotal declivity rather coarsely longitudinally rugose. Gaster smooth and shining.

Hairs yellow, not very abundant; clavate on thorax, pedicel and crown of head; short and erect on head and thorax, longer and somewhat reclinate on the pedicel and gaster. Hairs on antennae, legs and sides of head less conspicuous and appressed, except on the antennae, where they are suberect.

Rather dark reddish-brown, ventral portions of head, thorax and pedicel and the incisures of the gastric segments, yellowish. Legs and antennae yellowish, middle of femora and the antennal club darkbrown. Immature specimens have the thorax and pedicel more extensively yellow.

Queen

Length 4.5 mm. Mandibles densely striated and somewhat punctate. Clypeus with coarse longitudinal rugae, one of which forms a distinct carina in the middle of the sclerite. Frontal area opaque. Head with coarse and very regular longitudinal rugae, but little diverging behind and but slightly reticulate. On the upper surface of the head there are also a number of shallow but distinct foveolae in the interrugal furrows. Pronotum and pleurae coarsely longitudinally rugose; mesonotum shining, sparsely foveolate and rather indistinctly longitudinally rugose, especially in front. Scutellum and paraptera like the mesonotum, the former with indications of rugae only at its anterior border. Whole epinotum subopaque, coarsely rugose, even over the entire surface of the robust, pointed spines, which are as long as they are broad at their bases. Declivous surface of epinotum regularly transversely rugose. Petiole and postpetiole opaque, reticulate .and punctate-rugose, more coarsely on the sides than on the summits of the nodes; petiolar node in profile more acute than in the worker. Gaster very glabrous. Head, thorax and pedicel rich reddish-brown; gaster decidedly darker. Legs and antennre yellow. In the latter all the joints of the funiculus are distinctly longer than broad, and the club, which is not infuscated, is indistinct. Wing-insertions black, Hairs on the body sparse, yellow, not clavate like those of the worker, but more or less tapering.

Male

Length 2.5-3 mm. Mandibles dentate, overlapping with their blades. Clypeus convex, truncated in the middle in front. Antennae 13-jointed; scape slender, as long as joints 1-5 of the funiculus. First funicular joint fully twice as long as broad, much stouter than the succeeding joints, except those of the 4-jointed club; joints 3-7 longer than broad; three basal joints of club subequal, each not more than half as long as the terminal joint. Cheeks short. Thorax with very deep Mayrian and other sutures. Epinotum without indications of spines. Petiole and postpetiole longer and with lower. nodes than in the worker. Gaster of the usual shape.

Mandibles and clypeus somewhat shining, the latter with a few prominent and irregular longitudinal rugal. Head decidedly opaque, uniformly and densely punctate. Thorax shining, mesonotum, scutellum and pleural with faint, parallel, longitudinal striae. Pronotum and epinotum more reticulate-rugose. Petiole and postpetiole smooth and shining on the summits of the nodes, elsewhere subopaque, finely reticulate-rugose. Gaster smooth and shining.

Hairs covering the body sparse, whitish, non-clavate, longest and most conspicuous on the gaster, very small and appressed on the legs and antennal.

Black, thorax and pedicel more piceous, especially on their lateral and ventral surfaces. The following parts are yellowish, or yellow suffused with piceous: mandibles, except their teeth, which are black, antennae, legs and genitalia. Wings whitish-hyaline, veins and stigma colorless.

Type Material

Mackay (2000) USA. King's Canyon, Ormsby county, Nevada. Type series: American Museum of Natural History, Museum of Comparative Zoology [seen].

References

  • Alatorre-Bracamontes, C.E., Vásquez-Bolaños, M. 2010. Lista comentada de las hormigas (Hymenoptera: Formicidae) del norte de México. Dugesiana 17(1): 9-36.
  • Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 271, Combination in Temnothorax)
  • Cole, A. C., Jr. 1966. Ants of the Nevada Test Site. Brigham Young University Science Bulletin Biological Series. 7(3):1-27.
  • Fisher, B. L. 1997. A comparison of ant assemblages (Hymenoptera, Formicidae) on serpentine and non-serpentine soils in northern California. Insectes Sociaux. 44:23-33.
  • Johnson, R. A. and P. S. Ward. 2002. Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: a first overview. Journal of Biogeography. 29:1009-1026.
  • MacKay, W. P. 2000. A review of the New World ants of the subgenus Myrafant, (Genus Leptothorax) (Hymenoptera: Formicidae). Sociobiology. 36:265-444. (page 376, Senior synonym of eldoradensis, melanderi and rudis (unresolved junior homonym))
  • Mackay W. P. and Mackay, E. E. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
  • MontBlanc, E. M., J. C. Chambers, and P. F. Brussard. 2007. Variation in ant populations with elevation, tree cover, and fire in a pinyon-juniper-dominated watershed. Western North American Naturalist. 67:469-491.
  • Prebus, M.M. 2021. Taxonomic revision of the Temnothorax salvini clade (Hymenoptera: Formicidae), with a key to the clades of New World Temnothorax. PeerJ 9, e11514 (doi:10.7717/peerj.11514).
  • Ratchford, J. S., S. E. Wittman, E. S. Jules, A. M. Ellison, N. J. Gotelli, and N. J. Sanders. 2005. The effects of fire, local environment and time on ant assemblages in fens and forests. Diversity and Distributions. 11:487-497.
  • Smith, D. R. 1979. Superfamily Formicoidea. Pp. 1323-1467 in: Krombein, K. V., Hurd, P. D., Smith, D. R., Burks, B. D. (eds.) Catalog of Hymenoptera in America north of Mexico. Volume 2. Apocrita (Aculeata). Washington, D.C.: Smithsonian Institution Pr (page 1393, Combination in L. (Myrafant))
  • Snelling, R. R.; Borowiec, M. L.; Prebus, M. M. 2014. Studies on California ants: a review of the genus Temnothorax (Hymenoptera, Formicidae). ZooKeys 372:27-89. doi:10.3897/zookeys.372.6039
  • Ward, P. S. 2005. A synoptic review of the ants of California (Hymenoptera: Formicidae). Zootaxa. 936:1-68.
  • Wheeler, G. C. and J. Wheeler. 1978. Mountain ants of Nevada. Great Basin Naturalist. 38:379-396.
  • Wheeler, G. C. and J. Wheeler. 1986. The ants of Nevada. Natural History Museum of Los Angeles County, Los Angeles.
  • Wheeler, G. C.; Wheeler, J. 1973b. Ant larvae of four tribes: second supplement (Hymenoptera: Formicidae: Myrmicinae). Psyche (Camb.) 80: 70-82 (page 71, larva described)
  • Wheeler, W. M. 1903. A revision of the North American ants of the genus Leptothorax Mayr. Proceedings of the Academy of Natural Sciences of Philadelphia. 55:215-260.

References based on Global Ant Biodiversity Informatics

  • Adams T. A., W. J. Staubus, and W. M. Meyer. 2018. Fire impacts on ant assemblages in California sage scrub. Southwestern Entomologist 43(2): 323-334.
  • Allred D. M. 1982. Ants of Utah. The Great Basin Naturalist 42: 415-511.
  • Allred, D.M. 1982. The ants of Utah. Great Basin Naturalist 42:415-511.
  • Boulton A. M., Davies K. F. and Ward P. S. 2005. Species richness, abundance, and composition of ground-dwelling ants in northern California grasslands: role of plants, soil, and grazing. Environmental Entomology 34: 96-104
  • Clarke K.M., Fisher B.L. and LeBuhn G. 2008. The influece of urban park characteristics on ant (Hymenoptera, Formicidae) communities. Urban Ecosyst 11: 317-334
  • Cole A. C., Jr. 1934. An annotated list of the ants of the Snake River Plains, Idaho (Hymenoptera: Formicidae). Psyche (Cambridge) 41: 221-227.
  • Cole A. C., Jr. 1942. The ants of Utah. American Midland Naturalist 28: 358-388.
  • Cole, A.C. 1936. An annotated list of the ants of Idaho (Hymenoptera; Formicidae). Canadian Entomologist 68(2):34-39
  • Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
  • Des Lauriers J., and D. Ikeda. 2017. The ants (Hymenoptera: Formicidae) of the San Gabriel Mountains of Southern California, USA with an annotated list. In: Reynolds R. E. (Ed.) Desert Studies Symposium. California State University Desert Studies Consortium, 342 pp. Pages 264-277.
  • Fisher B. L. 1997. A comparison of ant assemblages (Hymenoptera, Formicidae) on serpentine and non-serpentine soils in northern California. Insectes Sociaux 44: 23-33
  • Fisher, B.L. 1997. A comparison of ant assemblages (Hymenoptera, Formicidae) on serpentine and non-serpentine soils in northern California. Insectes Sociaux 44:23-33.
  • Heron J. 2005. Ants of the South Okanagan Grasslands, British Columbia. Arthropods of Canadian Grasslands 11: 17-22.
  • Hoey-Chamberlain R. V., L. D. Hansen, J. H. Klotz and C. McNeeley. 2010. A survey of the ants of Washington and Surrounding areas in Idaho and Oregon focusing on disturbed sites (Hymenoptera: Formicidae). Sociobiology. 56: 195-207
  • Johnson, R.A. and P.S. Ward. 2002. Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: a first overview. Journal of Biogeography 29:1009–1026/
  • Knowlton G. F. 1970. Ants of Curlew Valley. Proceedings of the Utah Academy of Sciences, Arts and Letters 47(1): 208-212.
  • La Rivers I. 1968. A first listing of the ants of Nevada. Biological Society of Nevada, Occasional Papers 17: 1-12.
  • Mackay W. P. 2000. A review of the New World ants of the subgenus Myrafant, (genus Leptothorax) (Hymenoptera: Formicidae). Sociobiology 36: 265-444.
  • Mackay W. P. and Mackay, E. E. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
  • Mallis A. 1941. A list of the ants of California with notes on their habits and distribution. Bulletin of the Southern California Academy of Sciences 40: 61-100. 
  • MontBlanc E. M., J. C. Chambers, and P. F. Brussard. 2007. Variation in ant populations with elevation, tree cover, and fire in a Pinyon-Juniper-dominated watershed. Western North American Naturalist 67(4): 469–491.
  • Ostoja S. M., E. W. Schupp, and K. Sivy. 2009. Ant assemblages in intact big sagebrush and converted cheatgrass-dominates habitats in Tooele County, Utah. Western North American Naturalist 69(2): 223–234.
  • Parson G. L., G Cassis, A. R. Moldenke, J. D. Lattin, N. H. Anderson, J. C. Miller, P. Hammond, T. Schowalter. 1991. Invertebrates of the H.J. Andrews Experimental Forest, western Cascade Range, Oregon. V: An annotated list of insects and other arthropods. Gen. Tech. Rep. PNW-GTR-290. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 168 p.
  • Ratchford, J.S., S.E. Wittman, E.S. Jules, A.M. Ellison, N.J. Gotelli and N.J. Sanders. 2005. The effects of fire, local environment and time on ant assemblages in fens and forests. Diversity and Distributions 11:487-497.
  • Sanders, N.J. 2004.Immediate Effects of Fire on the Invasive Argentine Ant, Linepithema humile. The Southwestern Naturalist 49(2) :246-250
  • Snelling R.R., M. L. Borowiec, and M. M. Prebus. 2014. Studies on California ants: a review of the genus Temnothorax (Hymenoptera, Formicidae). ZooKeys 372: 27–89. doi: 10.3897/zookeys.372.6039
  • Staubus W. J., E. S. Boyd, T. A. Adams, D. M. Spear, M. M. Dipman, W. M. Meyer III. 2015. Ant communities in native sage scrub, non-native grassland, and suburban habitats in Los Angeles County, USA: conservation implications. Journal of Insect Conservervation 19:669–680
  • Ulyssea M. A., L. P. Prado, C. R. F. Brandao. 2015. Type specimens of the traditional Myrmicinae (Hymenoptera: Formicidae) ant tribes deposited in the Museu de Zoologia da Universidade de Sao Paulo, Brazil: Adelomyrmecini, Basicerotini, Blepharidattini, Crematogastrini, Formicoxenini, Lenomyrmecini, Myrmicini, Phalacromyrmecini, Pheidolini, Stegomyrmecini, Stenammini and Tetramoriini. Papeis Avulsos de Zoologia, Museu de Zoologia da Universidade de Sao Paulo 55(12): 175-204.
  • Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
  • Ward P. S. 2005. A synoptic review of the ants of California (Hymenoptera: Formicidae). Zootaxa 936: 1-68.
  • Wetterer, J.K., P.S. Ward, A.L. Wetterer, J.T. Longino, J.C. Trager and S.E. Miller. 2000. Ants (Hymenoptera:Formicidae) of Santa Cruz Island, California. Bulletin of the Southern California Academy of Science 99(1):25-31.
  • Wheeler G. C., and J. Wheeler. 1986. The ants of Nevada. Los Angeles: Natural History Museum of Los Angeles County, vii + 138 pp.
  • Wheeler W. M. 1903. A revision of the North American ants of the genus Leptothorax Mayr. Proceedings of the Academy of Natural Sciences of Philadelphia. 55: 215-260.
  • Wheeler W. M. 1917. The mountain ants of western North America. Proceedings of the American Academy of Arts and Sciences 52: 457-569.
  • Wheeler, G.C. and J. Wheeler. 1978. Mountain ants of Nevada. Great Basin Naturalist 35(4):379-396
  • Wheeler, G.C. and J. Wheeler. 1988. A checklist of the ants of Montana. Psyche 95:101-114
  • Wittman S. E., N. J. Sanders, A. M. Ellison, E. S. Jules, J. S. Ratchford, and N. J. Gotelli. Forthcoming. Species interactions and thermal constraints on ant community structure. Oikos 119.