Pogonomyrmex barbatus

Pogonomyrmex barbatus
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Pogonomyrmecini
Genus:	Pogonomyrmex
Species group:	barbatus
Species:	P. barbatus
Binomial name
	Pogonomyrmex barbatus; (Smith, F., 1858) Specimen Label
Synonyms
	Myrmica molefaciens Buckley, 1860; Pogonomyrmex barbatus nigrescens Wheeler, W.M., 1902;

Common Name

Red Harvester Ant
Language:	English

This species has large crater like mounds with a diameter of about 1 meter. They are cleared of vegetation and covered with gravel, if it is available. Brood and reproductives were found in the nests in July. They are extremely pugnacious and their sting is very painful. (Mackay and Mackay 2002)

Identification

This is usually an easily recognized species as it is large, red and aggressive. Occasionally it is difficult to distinguish specimens from those of Pogonomyrmex rugosus. The color usually works to distinguish this species (P. rugosus is dark with a con-trasting lighter gaster), but it may be necessary to examine the cephalic rugae to determine if they are fine or coarse as in P. rugosus. It can be separated from the other species, as it is much larger (total length over 8 mm, usually about 10 mm in total length). Workers from newly founded nests are small, and could be confused with Pogonomyrmex desertorum. The cephalic rugae are fine, but are not as fine as those of P. desertorum, which also has a shiny area at the posterior lateral corner, an area which is covered with rugae in P. barbatus. (Mackay and Mackay 2002)

Keys including this Species

Key to North American Pogonomyrmex

Distribution

Southwestern United States and northern Mexico.

Latitudinal Distribution Pattern

Latitudinal Range: 40.555794° to 16.66227°.


North Temperate	North Subtropical	Tropical	South Subtropical	South Temperate

Source: AntMaps

Distribution based on Regional Taxon Lists

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

Distribution based on AntMaps

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.

Habitat

Various habitats. Ranging from the edges of the desert and grasslands up to lower elevation pine forests, pinyon-juniper and oak forests, sagebrush and riparian habitats. It occurs in more mesic sites than P. rugosus.

Abundance

Quite common in northern Mexico.

Biology

An intermorphic queen has been described by Castano-Meneses & Vasquez-Bolanos (2007). These queens are characterized by less thoracic development, loss of wings, and absence of ocelli when compared to full queens. However, they have the same reproductive capacity in new colonies as do normal queens since they have the same number of ovarioles.

Genetics

The entire genome of P. barbatus (Smith et al. 2011) has been sequenced and was used for a study examining how parasitic ants with no worker caste may have altered their genome to arrive at a workerless state. In comparison to ants with a full complement of castes, there appeared to be no loss of genes in the parasitic ants. This suggests regulatory differences and not sequence differences predominate in gains and losses of castes (phenotypes). (Smith et al. 2015)

Palomeque et al. (2015) found class II mariner elements, a form of transposable elements, in the genome of this ant.

Ten Polymorphic microsatellite loci were identified by Volny, V. P. and Gordon, D. M., 2002.

Flight Period

			X	X	X	X	X	X
Jan	Feb	Mar	Apr	May	Jun	Jul	Aug	Sep	Oct	Nov	Dec

Source: antkeeping.info.

Check details at Worldwide Ant Nuptial Flights Data, AntNupTracker and AntKeeping.

Explore: Show all Flight Month data or Search these data. See also a list of all data tables or learn how data is managed.

Association with other Organisms

Other Ants

A host species for the workerless social parasite Pogonomyrmex anergismus.

This species is a xenobiont for the ant Monomorium minimum (a xenobiont) in United States (Kanizsai et al., 2013; Wheeler, 1901).

Coleoptera

This species is prey for the tiger beetles Cicindela formosa gibsoni, Cicindela pugetana, Cicindela pulchra, Cicindela scutellaris and Cicindela tranquebarica kirbyi (predators) in the United States (Valenti & Gaimari, 2000; Polidori et al., 2020).

Orthoptera

This species is a host for the cricket Myrmecophilus nebrascensis (a myrmecophile) in United States.

Fungi

This species is a host for the fungus Myrmicinosporidium durum (a pathogen) in Texas, United States (Sanchez-Peña et al., 1993; Espadaler & Santamaria, 2012).

Nematodes

Pogonomyrmex barbatus is an intermediate host for the nematode Skrjabinoptera phrynosoma, which lives in the stomach of the Texas horned lizard, Phrynosoma cornutum (Lee, 1957).

Castes

Worker

Images from AntWeb


Worker. Specimen code casent0102894. Photographer Jen Fogarty, uploaded by California Academy of Sciences.	Owned by CAS, San Francisco, CA, USA.

Images from AntWeb


Worker. Specimen code casent0006306. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by CAS, San Francisco, CA, USA.

Nomenclature

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

barbatus. Myrmica barbata Smith, F. 1858b: 130 (q.) MEXICO. Mayr, 1870b: 971 (w.); Mayr, 1887: 610 (m.); Wheeler, W.M. 1900b: 20 (l.); Taber, Cokendolpher & Francke, 1988: 51 (k.). Combination in Pogonomyrmex: Mayr, 1868b: 170. Senior synonym of molefaciens: Mayr, 1886c: 365; Creighton, 1950a: 119; Cole, 1968: 56; of nigrescens: Cole, 1968: 56. See also: Wheeler, G.C. & Wheeler, J. 1953a: 107; Petralia & Vinson, 1980: 381; Smith, D.R. 1979: 1353.
molefaciens. Myrmica (Atta) molefaciens Buckley, 1860b: 445 (w.q.) U.S.A. Emery, 1895c: 308 (m.). Combination in Pogonomyrmex: Mayr, 1886c: 365. Subspecies of barbatus: Forel, 1886b: xlii; Emery, 1895c: 308; Forel, 1901c: 128. Junior synonym of barbatus: Mayr, 1886c: 365; Creighton, 1950a: 119; Cole, 1968: 56.
nigrescens. Pogonomyrmex barbatus var. nigrescens Wheeler, W.M. 1902c: 389 (w.) U.S.A. Junior synonym of fuscatus: Creighton, 1950a: 119; of barbatus: Cole, 1968: 56.

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

Description

Worker

Cole (1968) - HL 1.98-2.43 mm, HW 2.05-2.70 mm, CI 103.6-111.1, SL 1.33-1.71 mm, SI 63.3-68.4, EL 0.42-0.53 mm, EW 0.23-0.34 mm, OI 21.2-21.8, WL 2.24-2.89 mm, PNL 0.46-0.65 mm, PNW 0.49-0.68 mm, PPL 0.42-0.65 mm, PPW 0.34-0.91 mm.

Mandible as shown in Pl. III, Fig. 17; teeth short, robust, blunt; apical and subapical teeth generally longer than others; ultimate basal tooth longer and broader than penultimate, slightly but distinctly offset from the moderately convex basal margin.

Base of antennal scape like that of rugosus (Pl. IV, Fig. 15), moderately strongly curved in proximal one-half its length, somewhat constricted but not flattened along the bend. Basal enlargement broad and strong; apex of superior lobe varies from blunt and subtruncate to subacute; gradient of superior declivity variable, but generally rather steep and forming a strong angle with the flange; lip strong and hook. shaped, its margin thin and directed distad and strongly inward forming a prominent concavity; point present or absent, when present varying from weak to strong; basal flange narrow, straight, extending from lip to apex of superior lobe; lateral surface of basal enlargement strongly, evenly, broadly, and longitudinally impressed, forming a prominent, moderately deep, elongate, saucerlike depression, the lower margin of which is bordered by the longitudinal peripheral carina (when present), the proximal extension of which forms the point.

Contour of thorax, petiole, and postpetiole as portrayed in Pl. V, Fig. 1. Epinotal spines always present, varying from short to long. Petiole and postpetiole in dorsal view as shown in Pl. VII, Fig. 17.

Cephalic rugae moderately to very fine, rather even, moderately to closely spaced; interrugal spaces shining to subopaque, without sculpture, or with fine punctures and/or foveae, often with granulations and sometimes with a few broken rugulae. Thoracic rugae fine, rather even, and closely set to moderately coarse and irregular; often strongest on pronotum, where they may be wavy but do not form strong reticulations with deep interrugal spaces. Interrugal spaces of thorax shining to subopaque, with or without punctures. Rugae on petiolar and postpetiolar nodes moderately fine to coarse, rather even, not forming coarse reticulations.

Body color varying from a concolorous light ferrugineous to very deep red.

Queen

Cole (1968) - HL 2.13-2.17 mm, HW 2.93-3.01 mm, CI 118.6-125.1, SL 1.63-1.67 mm, SI 54.9-55.6, EL 0.57-0.61 mm, EW 0.30-0.38 mm, OI 23.1-25.1, WL 3.80-3.81 mm, PNL 0.68-0.76 mm, PNW 0.68-0.84 mm, PPL 0.61-0.68 mm, PPW 0.95-0.99 mm.

Essentially similar to the worker, but with the usual female characters. Cephalic and scutal rugae rather fine, closely spaced. Scutellum smooth, very shining, longitudinal striae weak or absent. Epinotal spines short, prominent, robust, sharp. Dorsum of petiolar node rugose, the rugae not forming reticulations. Body color light to deep ferrugineous red.

Male

Cole (1968) - HL 1.60-2.09 mm, HW 2.09-2.36 mm, CI 112.9-130.6, SL 0.84-0.87 mm, SI 35.5-41.6, EL 0.61-0.61 mm, EW 0.42-0.46 mm, OI 29.2-29.2, WL 3.46-3.57 mm, PNL 0.49-0.53 mm, PNW 0.84-0.99 mm, PPL 0.57-0.65 mm, PPW 1.10-1.18 mm.

Mandible as shown in Pl. VIII, Fig. 11. Outer surface of base of antennal scape strongly flattened or broadly concave. Paramere as illustrated in Pl. X, Fig. 10 and Pl. XI, Fig. 11; inner dorsal bonier with a deep, rather broad emargination which strongly separates the terminal lobe from the base. Body gcnerally a concolorous clear yellow.

Karyotype

See additional details at the Ant Chromosome Database.

Explore: Show all Karyotype data or Search these data. See also a list of all data tables or learn how data is managed.

n = 16, 2n = 32 (USA) (Taber et al., 1988).

References

Borowiec, M.L. 2019. Convergent evolution of the army ant syndrome and congruence in big-data phylogenetics. Systematic Biology 68, 642–656 (doi:10.1093/sysbio/syy088).
Branstetter, M.G., Danforth, B.N., Pitts, J.P., Faircloth, B.C., Ward, P.S., Buffington, M.L., Gates, M.W., Kula, R.R., Brady, S.G. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27, 1019–1025 (doi:10.1016/j.cub.2017.03.027).
Bulter, I. 2020. Hybridization in ants. Ph.D. thesis, Rockefeller University.
Burford, B.P., Lee, G., Friedman, D.A., Brachmann, E., Khan, R., MacArthur-Waltz, D.J., McCarty, A.D., Gordon, D.M. 2018. Foraging behavior and locomotion of the invasive Argentine ant from winter aggregations. PLOS ONE 13, e0202117 (doi:10.1371/JOURNAL.PONE.0202117).
Cantone S. 2017. Winged Ants, The Male, Dichotomous key to genera of winged male ants in the World, Behavioral ecology of mating flight (self-published).
Castano-Meneses, G., Vásquez-Bolaños, M. 2007. Intermorphic queen in the harvester ant Pogonomyrmex barbatus (F. Smith) (Hymenoptera: Formicidae) in Tonala, Jalisco, Mexico. Southwestern Entomologist 32: 131-133.
Cole, A. C., Jr. 1968. Pogonomyrmex harvester ants. A study of the genus in North America. Knoxville, Tenn.: University of Tennessee Press, x + 222 pp. (page 56, Senior synonym of molefaciens, Senior synonym of nigrescens)
Creighton, W. S. 1950a. The ants of North America. Bulletin of the Museum of Comparative Zoology 104: 1-585 (page 119, Senior synonym of molefaciens)
Cushing, P.E. 2012. Spider-ant associations: An updated review of myrmecomorphy, myrmecophily, and myrmecophagy in spiders. Psyche: A Journal of Entomology 2012, 1–23 (doi:10.1155/2012/151989).
Darras, H., Kuhn, A., Aron, S. 2019. Evolution of hybridogenetic lineages in Cataglyphis ants. Molecular Ecology 28, 3073–3088 (doi:10.1111/mec.15116).
Del Toro, I., Robbons, R.R., Pelini, S.L. 2012. The little things that run the world revisited: a review of ant-mediated ecosystem services and disservices (Hymenoptera: Formicidae). Myrmecological News 17: 133-146.
Ellison, A.M., Gotelli, N.J. 2021. Ants (Hymenoptera: Formicidae) and humans: from inspiration and metaphor to 21st-century symbiont. Myrmecological News 31: 225-240 (doi:10.25849/MYRMECOL.NEWS_031:225).
Espadaler, X., Santamaria, S. 2012. Ecto- and Endoparasitic Fungi on Ants from the Holarctic Region. Psyche Article ID 168478, 10 pages (doi:10.1155/2012/168478).
Gordon, D.M. 1988. Nest-plugging: interference competition in desert harvester ants (Novomessor cockerelli and Pogonomyrmex barbatus). Oecologia 75: 114-118. (doi:10.1007/BF00378823).
Hoey-Chamberlain, R., Rust, M.K., Klotz, J.H. 2013. A review of the biology, ecology and behavior of Velvety Tree Ants of North America. Sociobiology 60(1): 1-10.
Hoey-Chamberlain, R.V. 2012. Food preference, survivorship, and intraspecific interactions of Velvety Tree Ants. M.S. thesis, University of California, Riverside.
Horna-Lowell, E., Neumann, K.M., O’Fallon, S., Rubio, A., Pinter-Wollman, N. 2021. Personality of ant colonies (Hymenoptera: Formicidae) – underlying mechanisms and ecological consequences. Myrmecological News 31: 47-59 (doi:10.25849/MYRMECOL.NEWS_031:047).
Ivanov, K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87 (doi:10.3897@jhr.70.35207).
Jacobs, S. 2020. Population genetic and behavioral aspects of male mating monopolies in Cardiocondyla venustula (Ph.D. thesis).
Kanizsai, O., Lőrinczi, G., Gallé, L. 2013. Nesting associations without interdependence: A preliminary review on plesiobiosis in ants. Psyche 2013, 238602 (doi:10.1155/2013/238602).
Laciny, A. 2021. Among the shapeshifters: parasite-induced morphologies in ants (Hymenoptera, Formicidae) and their relevance within the EcoEvoDevo framework. EvoDevo 12, 2 (doi:10.1186/s13227-021-00173-2).
Lau, M.K., Ellison, A.M., Nguyen, A., Penick, C., DeMarco, B., Gotelli, N.J., Sanders, N.J., Dunn, R.R., Helms Cahan, S. 2019. Draft Aphaenogaster genomes expand our view of ant genome size variation across climate gradients. PeerJ 7, e6447 (doi:10.7717/PEERJ.6447).
Lee, S.H. 1957. The life cycle of Skrjabinoptera phrynosoma (Ortlepp) Schulz, 1927 (Nematoda: Spiruroidea), a gastric nematode of Texas horned toads, Phrynosoma cornutum. The Journal of Parasitology 43: 66–75.
Lo, N., Beekman, M., Oldroyd, B.P. 2019. Caste in social insects: Genetic influences over caste determination. In: Encyclopedia of Animal Behavior (Second Edition), pp. 274–281 (doi:10.1016/B978-0-12-809633-8.20759-0).
Maák, I., Roelandt, G., d'Ettorre, P. 2020. A small number of workers with specific personality traits perform tool use in ants. eLife 9, e61298 (doi:10.7554/elife.61298).
Mackay, W. P. and E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Edwin Mellen Press, Lewiston, NY.
Mayr, G. 1868b. Formicidae novae Americanae collectae a Prof. P. de Strobel. Annu. Soc. Nat. Mat. Modena 3: 161-178 (page 170, Combination in Pogonomyrmex)
Mayr, G. 1870b. Neue Formiciden. Verh. K-K. Zool.-Bot. Ges. Wien 20: 939-996 (page 971, worker described)
Mayr, G. 1886c. Notizen über die Formiciden-Sammlung des British Museum in London. Verh. K-K. Zool.-Bot. Ges. Wien 36: 353-368 (page 365, Senior synonym of molefaciens)
Mayr, G. 1887. Südamerikanische Formiciden. Verh. K-K. Zool.-Bot. Ges. Wien 37: 511-632 (page 610, male described)
Möglich, M. 1978. Social organization of nest emigration in Leptothorax (Hym., Form.). Insectes Sociaux 25(3), 205–225 (doi:10.1007/bf02224742).
Palomeque, T., O. Sanllorente, X. Maside, J. Vela, P. Mora, M. I. Torres, G. Periquet, and P. Lorite. 2015. Evolutionary history of the Azteca-like mariner transposons and their host ants. Science of Nature. 102. doi:10.1007/s00114-015-1294-3
Parker, J.D., Rissing, S.W. 2002. Molecular evidence for the origin of workerless social parasites in the ant genus Pogonomyrmex. Evolution 56: 2017-2028.
Perfilieva, K.S. 2023. Cretaceous-Burmese-amber ants: Morphological features and community structure. Biology Bulletin Reviews 131, 38–54 (doi:10.1134/s207908642301005x).
Petralia, R. S.; Vinson, S. B. 1980 [1979]. Comparative anatomy of the ventral region of ant larvae, and its relation to feeding behavior. Psyche (Camb.) 86: 375-394 (page 381, see also)
Pirk, G.I., Casenave, J.L.de 2006. Diet and seed removal rates by the harvester ants Pogonomyrmex rastratus and Pogonomyrmex pronotalis in the central Monte desert, Argentina. Insectes Sociaux 53, 119–125 (doi:10.1007/s00040-005-0845-6).
Plowes, N.J.R., Johnson, R.A., Holldobler, B. 2013. Foraging behavior in the ant genus Messor (Hymenoptera: Formicidae: Myrmicinae). Myrmecological News 18, 33-49.
Polidori, C., Rodriguez-Flores, P.C., Garcia-Paris, M. 2020. Ants as prey for the endemic and endangered Spanish tiger beetle Cephalota dulcinea (Coleoptera: Carabidae). Annales de la Société entomologique de France (N.S.) (doi:10.1080/00379271.2020.1791252).
Roux, J., Privman, E., Moretti, S., Daub, J.T., Robinson-Rechavi, M., Keller, L. 2014. Patterns of positive selection in seven ant genomes. Molecular Biology and Evolution 31, 1661–1685 (doi:10.1093/molbev/msu141).
Ruano, F., Tinaut, A., Soler, J.J. 2000. High surface temperatures select for individual foraging in ants. Behavioral Ecology 11, 396-404.
Sanchez-Peña, S.R., Buschinger, A., Humber, R.A. 1993. Myrmicinosporidium durum, an enigmatic fungal parasite of ants. Journal of Invertebrate Pathology 61: 90-96 (doi:10.1006/jipa.1993.1016).
Smith C.R., Smith C.D., Robertson H.M., Helmkampf M., Zimin A., Yandell M., Holt C., Hu H., Abouheif E., and R. Benton. 2011. Draft genome of the red harvester ant Pogonomyrmex barbatus. Proc Natl Acad Sci. 108:5667–5672.
Smith, C. R. et al. 2011. Draft genome of the red harvester ant Pogonomyrmex barbatus. PNAS. 108(14):5667-5672. doi:10.1073/pnas.1007901108
Smith, C. R., S. H. Cahan, C. Kemena, S. G. Brady, W. Yang, E. Bornberg-Bauer, T. Eriksson, J. Gadau, M. Helmkampf, D. Gotzek, M. O. Miyakawa, A. V. Suarez, and A. Mikheyev. 2015. How Do Genomes Create Novel Phenotypes? Insights from the Loss of the Worker Caste in Ant Social Parasites. Molecular Biology and Evolution. 32:2919-2931. doi:10.1093/molbev/msv165
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 1353, see also)
Smith, F. 1858b. Catalogue of hymenopterous insects in the collection of the British Museum. Part VI. Formicidae. London: British Museum, 216 pp. (page 130, queen described)
Stukalyuk, S., Radchenko, Y., Gonchar, O., Akhmedov, A., Stelia, V., Reshetov, A., Shymanskyi, A. 2021. Mixed colonies of Lasius umbratus and Lasius fuliginosus (Hymenoptera, Formicidae): when superparasitism may potentially develop into coexistence: a case study in Ukraine and Moldova. Halteres 12, 25–48 (doi:10.5281/zenodo.5753121).
Taber, S. W.; Cokendolpher, J. C.; Francke, O. F. 1988. Karyological study of North American Pogonomyrmex (Hymenoptera: Formicidae). Insectes Soc. 35: 47-60 (page 51, karyotype described)
Ulysséa, M.A., Farder-Gomes, C.F., Prado, L.P. 2024. Biological notes, nest architecture, and morphology of the remarkable ant Hylomyrma primavesi Ulysséa, 2021 (Hymenoptera: Formicidae: Myrmicinae). Myrmecological News 34, 1-20 (doi:10.25849/MYRMECOL.NEWS_034:001).
Valenti, M.A., Gaimari, S.D. 2000. Of tiger beetles and ubiquitous ants: hazardous living amongst tenacious prey. Cicindela 32: 19–29.
Varela-Hernández, F., Medel-Zosayas, B., Martínez-Luque, E.O., Jones, R.W., De la Mora, A. 2020. Biodiversity in central Mexico: Assessment of ants in a convergent region. Southwestern Entomologist 454: 673-686.
Volny, V. P. and D. M. Gordon. 2002. Characterization of polymorphic microsatellite loci in the red harvester ant, Pogonomyrmex barbatus. Molecular Ecology Notes. 2(3):302-303. doi:10.1046/j.1471-8286.2002.00243.x
Walsh, J., Pontieri, L., d'Ettorre, P., Linksvayer, T.A. 2020. Ant cuticular hydrocarbons are heritable and associated with variation in colony productivity. Proceedings of the Royal Society B: Biological Sciences 287: 20201029 (doi:10.1098/RSPB.2020.1029).
Wang, C., Lin, C.-C., Keller, R.A., Billen, J. 2021. The ‘hairwheels’ in Strumigenys ants are not glandular. Asian Myrmecology 13: e013004 (doi:10.20362/am.013004).
Wheeler, G. C.; Wheeler, J. 1953a [1952]. The ant larvae of the myrmicine tribe Myrmicini. Psyche (Camb.) 59: 105-125 (page 107, see also)
Wheeler, W. M. 1900b. A study of some Texan Ponerinae. Biol. Bull. (Woods Hole) 2: 1-31 (page 20, larva described)
Wheeler, W.M. 1901. The compound and mixed nests of American ants. The American Naturalist 35(2-3), 513–539, 791–818.
Zhou, X., Slone, J.D., Rokas, A., Berger, S.L., Liebig, J., Ray, A., Reinberg, D., Zwiebel, L.J. 2012. Phylogenetic and transcriptomic analysis of chemosensory receptors in a pair of divergent ant species reveals sex-specific signatures of odor coding. PLoS Genetics 8, e1002930 (doi:10.1371/journal.pgen.1002930).

References based on Global Ant Biodiversity Informatics

Alatorre-Bracamontes, C.E. and M Vasquez-Bolanos. 2010. Lista comentada de las hormigas (Hymenoptera: Formicidae) del norte de México. Dugesiana 17(1):9-36
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.
Andersen A. N. 1997. Functional Groups and Patterns of Organization in North American Ant Communities: A Comparison with Australia. Journal of Biogeography. 24: 433-460
Anderson K. E., L. Zeltzer, R. P. Overson, and W. H. Clark. 2010. Identification of cryptic hosts for two inquiline parasites of the seed-harvester ant Pogonomyrmex and new localities for P. anergismus (Hymenoptera: Formicidae). The Southwestern Naturalist 55: 532-538.
Author Unknown. 2004. Mirmecofauna (Hymenoptera: Formicidae) comu?n del estero El Salado y Puerto Vallarta, Jalisco, Me?xico. Dugesiana 11(1):13-20.
Castaño-Meneses G., and M. Vásquez-Bolaños. 2007. Intermorphic queen in the harvester ant Pogonomyrmex barbatus (F. Smith) (Hymenoptera: Formicidae) in Tonalá, Jalisco, México. Southwestern Entomologist, 32 (2): 131-133
Ceballos G., R. List, G. Garduno, R. Lopez Cano, M. J. Munozcano Quintanar, E. Collado, and J. E. San Roman. 2009. La diversidad biologica del Estado de Mexico. Biblioteca Mexiquense del Bicentenario 530 pages.
Clark, R.M., K.E. Anderson, J. Gadau and J.H. Fewell. 2006. Behavioral Regulation of Genetic Caste Determination in a Pogonomyrmex Population with Dependent Lineages. Ecology 87:2201-2206.
Cokendolpher J.C., Reddell J.R., Taylor S.J, Krejca J.K., Suarez A.V. and Pekins C.E. 2009. Further ants (Hymenoptera: Formicidae) from caves of Texas [Hormigas (Hymenoptera: Formicdae) adicionales de cuevas de Texas]. Texas Memorial Museum Speleological Monographs, 7. Studies on the cave and endogean fauna of North America, V. Pp. 151-168
Cole A. C., Jr. 1937. An annotated list of the ants of Arizona (Hym.: Formicidae). [concl.]. Entomological News 48: 134-140.
Cole A. C., Jr. 1954. Studies of New Mexico ants. VII. The genus Pogonomyrmex with synonymy and a description of a new species (Hymenoptera: Formicidae). Journal of the Tennessee Academy of Science 29: 115-121.
Cook J. L. 2003. Conservation of biodiversity in an area impacted by the red imported fire ant, Solenopsis invicta (Hymenoptera: Formicidae). Biodiversity and Conservation 12: 187195.
Coronado-Blanco J. M., D. A. Dubovikoff, E. Ruiz-Cancino, M. Vasquez-Bolanos, K. Y. Flores-Maldonado, and J. V. Horta-Vega. 2013. Formicidae (Hymenoptera) del estado de Tamaulipas, Mexico. CienciaUat 25(1): 12-17.
Cover S. P., and R. A. Johnson. 20011. Checklist of Arizona Ants. Downloaded on January 7th at http://www.asu.edu/clas/sirgtools/AZants-2011%20updatev2.pdf
Dash S. T. and L. M. Hooper-Bui. 2008. Species diversity of ants (Hymenoptera: Formicidae) in Louisiana. Conservation Biology and Biodiversity. 101: 1056-1066
Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
DuBois M. B. 1981. New records of ants in Kansas, III. State Biological Survey of Kansas. Technical Publications 10: 32-44
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1111
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1112
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1113
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1114
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1115
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1116
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1117
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1118
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1119
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1120
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1121
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1122
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1123
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1124
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1125
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1126
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1127
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1128
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1129
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1130
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1131
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1132
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1133
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1134
DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1135
Fernandes, P.R. XXXX. Los hormigas del suelo en Mexico: Diversidad, distribucion e importancia (Hymenoptera: Formicidae).
Gregg, R.T. 1963. The Ants of Colorado.
Guzman-Mendoza R., G. Castano-Meneses, and M. de Carmen Hererra-Fuentes. 2010. Spatial and temporal variation of the diversity ants in the Botanic Garden from Zapotitlán de las Salinas Valley, Puebla. Revista Mexicana de Biodiversidad 81: 427- 435.
Hernandez, F. Varela and G. Castano-Meneses. 2010. Checklist, Biological Notes and Distribution of Ants (Hymenoptera: Formicidae) from Barranca de Metztitlán Biosphere Reserve, Hidalgo, Mexico. Sociobiology 56(2):397-434
Hernandez-Flores, J., M. Osorio-Beristain, and C. Martinez-Garza, 2016. Ant foraging as an indicator of Tropical Dry Forest restoration. Environmental Entomology 45: 991-994.
Herrera-Fuentes M. C., A. Navarrete-Jimenez, J. A. Zavala-Hurtado, J. Orendain-Mendez, and J. Campos-Serrano. 2015. Diversity of ants of the botanical garden from Zapotitlan, Puebla. Entomología Mexicana 2: 558-563.
Hess C. G. 1958. The ants of Dallas County, Texas, and their nesting sites; with particular reference to soil texture as an ecological factor. Field and Laboratory 26: 3-72.
Ivanov K., L. Hightower, S. T. Dash, and J. B. Keiper. 2019. 150 years in the making: first comprehensive list of the ants (Hymenoptera: Formicidae) of Virginia, USA. Zootaxa 4554 (2): 532–560.
Johnson R. A., and C. S. Moreau. 2016. A new ant genus from southern Argentina and southern Chile, Patagonomyrmex (Hymenoptera: Formicidae). Zootaxa 4139: 1-31.
Johnson R. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
Johnson, R.A. 2000. Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research 9(2):377-384.
Johnson, R.A. 2002. Semi-Claustral Colony Founding in the Seed-Harvester Ant Pogonomyrmex californicus: A Comparative Analysis of Colony Founding Strategies. Oecologia 132(1):60-67
Kempf, W.W. 1972. Catalago abreviado das formigas da regiao Neotropical (Hym. Formicidae) Studia Entomologica 15(1-4).
Kusnezov N. 1951. El género Pogonomyrmex Mayr (Hym., Formicidae). Acta Zoologica Lilloana 11: 227-333.
LeBrun E. G., R. M. Plowes, and L. E. Gilbert. 2015. Imported fire ants near the edge of their range: disturbance and moisture determine prevalence and impact of an invasive social insect. Journal of Animal Ecology,81: 884–895.
Lopez, A. S., M. Vasquez-Bolanos, and G. A. Q. Rocha. 2015. Hormigas (Hymenoptera: Formicidae) del Cerro de la Culebra, Arandas, Jalisco, Mexico. Dugesiana 19: 151-155.
Luna, P., J. H. Garcia-Chavez, and W. Dattilo. 2018. Complex foraging ecology of the red harvester ant and its effect on the soil seed bank. Acta Oecologica 86 57-65.
Luna, P., Y. Penaloza-Arellanes, A. L. Castillo-Meza, J. H. Garcia-Chavez, and W. Dattilo. 2018. Beta diversity of ant-plant interactions over day-night periods and plant physiognomies in a semiarid environment. Journal of Arid Environments 156: 69-76.
MacGown J. A., T. L. Schiefer, and M. G. Branstetter. 2015. First record of the genus Leptanilloides (Hymenoptera: Formicidae: Dorylinae) from the United States. Zootaxa 4006 (2): 392–400.
MacKay W. P., and S. Van Vactor. 1985. New host record for the social parasite Pogonomyrmex anergismus (Hymenoptera: Formicidae). Proceedings of the Entomological Society of Washington 87: 863.
Mackay, W.P., E.E. Mackay, J.F. Perez Dominguez, L.I. Valdez Sanchez and P.V. Orozco. 1985. Las hormigas del estado de Chihuahua Mexico: El genero Pogonomyrmex (Hymenoptera: Formicidae) . Sociobiology 11(1):39-54
Mackay W. P., and E. E. Mackay. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Lewiston, New York: Edwin Mellen Press, 400 pp.
Mackay, W., D. Lowrie, A. Fisher, E. Mackay, F. Barnes and D. Lowrie. 1988. The ants of Los Alamos County, New Mexico (Hymenoptera: Formicidae). pages 79-131 in J.C. Trager, editor, Advances in Myrmecololgy.
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.
McDonald D. L., D. R. Hoffpauir, and J. L. Cook. 2016. Survey yields seven new Texas county records and documents further spread of Red Imported Fire Ant, Solenopsis invicta Buren. Southwestern Entomologist, 41(4): 913-920.
Michigan State University, The Albert J. Cook Arthropod Research Collection. Accessed on January 7th 2014 at http://www.arc.ent.msu.edu:8080/collection/index.jsp
Moody J. V., and O. F. Francke. 1982. The Ants (Hymenoptera, Formicidae) of Western Texas Part 1: Subfamily Myrmicinae. Graduate Studies Texas Tech University 27: 80 pp.
Morrison, L.W. 2002. Long-Term Impacts of an Arthropod-Community Invasion by the Imported Fire Ant, Solenopsis invicta. Ecology 83(8):2337-2345
Mott, B. M., J. Gadau, and K. E. Anderson. 2015. Phylogeography of Pogonomyrmex barbatus and P. rugosus harvester ants with genetic and environmental caste determination. Ecology and Evolution 5: 2798-2826.
O'Keefe S. T., J. L. Cook, T. Dudek, D. F. Wunneburger, M. D. Guzman, R. N. Coulson, and S. B. Vinson. 2000. The Distribution of Texas Ants. The Southwestern Entomologist 22: 1-92.
Olsen O. W. 1934. Notes on the North American harvesting ants of the genus Pogonomyrmex Mayr. Bulletin of the Museum of Comparative Zoology 77: 493-514.
Parker, J.D. and S.W. Rissing. 2002. Molecular Evidence for the Origin of Workerless Social Parasites in the Ant Genus Pogonomyrmex. Evolution 56(10):2017-2028
Pergande, T. 1895. Mexican Formicidae. Proceedings of the California Academy of Sciences Ser. 2 :850-896
Quintana Ascencio, P. F., and M. Gonzalez Espinosa. 1990. Variacion estacional en la dieta de Pogonomyrmex barbatus (Hymenoptera: Formicidae) en nopaleras del centro de Mexico. Folia Entomologica Mexicana 80: 245-261.
Rico-Gray,V., J.G. Garcia-Franco, M. Palacios-Rios, C. Diaz-Castelazo, V. Parra-Tabla and J.A. Navarro. 1998. Geographical and Seasonal Variation in the Richness of Ant-Plant Interactions in Mexico. Biotropica 30(2):190-200.
Rios-Casanova, L., A. Valiente-Banuet, and V. Rico-Gray. (2004). Las hormigas del Valle de Tehuacan (Hymenoptera: Formicidae): una comparacion con otras zonas aridas de Mexico. Acta Zoologica Mexicana 20: 37-54.
Rivas-Arancibia, S. P., H. Carrillo-Ruiz, A. Bonilla-Arce, D. M. Figueroa-Castro, and A. R. Andres-Hernandez. 2014. Effect of disturbance on the ant community in a semiarid region of central Mexico. Applied Ecology and Environmental Research 12: 703-716.
Roeder K. A., and D. V. Roeder. 2016. A checklist and assemblage comparison of ants (Hymenoptera: Formicidae) from the Wichita Mountains Wildlife Refuge in Oklahoma. Check List 12(4): 1935.
Ríos-Casanova, L., A. Valiente-Banuet and V. Rico-Gray. 2004. Las hormigas del Valle de Tehuacan (Hymenoptera: Formicidae): Una comparacion con otras zonas aridas de Mexico. Acta Zoologica Mexicana 20(1):37-54
Serrano-Cardozo, V. H., J. A. Lemos-Espinal, and G. R. Smith. 2008. Comparative diet of three sympatric Sceloporus in the semiarid Zapotitlan Valley, Mexico. Revista Mexicana de Biodiversidad 79: 427-434.
Smith M. R. 1935. A list of the ants of Oklahoma (Hymen.: Formicidae). Entomological News 46: 235-241.
Smith M. R. 1936. A list of the ants of Texas. Journal of the New York Entomological Society 44: 155-170.
Taber S. W., J. C. Cokendolpher, and O. F. Francke. 1988. Karyological study of North American Pogonomyrmex (Hymenoptera: Formicidae). Insectes Soc. 35: 47-60.
Van Pelt, A. 1983. Ants of the Chisos Mountains, Texas (Hymenoptera: Formicidae) . Southwestern Naturalist 28:137-142.
Varela-Hernandez, F., M. Rocha-Ortega, R. W. Jones, and W. P. Mackay. 2016. Insectos: Hormigas (Formicidae) del estado de Queretaro, Mexico. Pages 397-404 in W. Jones., and V. Serrano-Cardenas, editors. Historia Natural de Queretaro. Universidad Autonoma de Queretaro, Mexico.
Varela-Hernandez, F., M. Rocha-Ortega, W. P. Mackay, and R. W. Jones. 2016. Lista preliminar de las hormigas (Hymenoptera: Formicidae) del estado de Queretaro, Mexico. Pages 429-435 in . W. Jones., and V. Serrano-Cardenas, editors. Historia Natural de Queretaro. Universidad Autonoma de Queretaro, Mexico.
Vasquez Bolanos M., and J. Escoto Rocha. 2018. Ants (Hymenoptera: Formicidae) of Aguascalientes. Investigacion y Ciencia 24(68): 36-40.
Vasquez-Bolanos M. 2011. Checklist of the ants (Hymenoptera: Formicidae) from Mexico. Dugesiana 18(1): 95-133.
Vasquez-Bolanos M., and J. L. Navarrete-Heredia. 2004. Checklist of the ants (Hymenoptera: Formicidae) from Jalisco State, Mexico. Sociobiology 43(2): 351-365
Villalvazo-Palacios, M., and J. F. Perez-Dominguez . 2011. Diversidad de hormigas (Insecta: Hymenoptera: Formicidae) del Area Natural Protegida Sierra de Quila Jalisco, Mexico. Pages 66-73 in Villavicencio, G. R., P. A. L. Santiago, E. V. C. Rosas, and L. L. Hernandez, editors. I Foro de conocimiento, uso y gestion del area Natural Protegida Sierra de Quila. Memorias. Organica Editores. Mexico.
Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
Warren, L.O. and E.P. Rouse. 1969. The Ants of Arkansas. Bulletin of the Agricultural Experiment Station 742:1-67
Warriner M. D. 2011. Status of the harvester ant (Pogonomyrmex spp.; Hymenoptera: Formicidae) fauna of Arkansas. Journal of the Kansas Entomological Society 84(2):160-162.
Wheeler G. C., and J. Wheeler J. 1989. A checklist of the ants of Oklahoma. Prairie Naturalist 21: 203-210.
Wheeler W. M. 1901. Notices biologiques sur les fourmis Mexicaines. Annales de la Société Entomologique de Belgique. 45: 199-205.
Wheeler W. M. 1901. The compound and mixed nests of American ants. Part II. The known cases of social symbiosis among American ants. American Naturalist. 35: 513-539.
Wheeler W. M. 1902. New agricultural ants from Texas. Psyche (Cambridge). 9: 387-393.
Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
Young J., and D. E. Howell. 1964. Ants of Oklahoma. Miscellaneous Publication. Oklahoma Agricultural Experimental Station 71: 1-42.
Young, J. and D.E. Howell. 1964. Ants of Oklahoma. Miscellaneous Publications of Oklahoma State University MP-71