Pheidole pilifera

The last published taxonomic treatment of this species (Wilson 2003) hypothesized this is a morphologically variable, broadly distributed species that is found in a wide range of habitats. Wilson's treatment and earlier work by Cole (1950) synonymized numerous subspecies (see the bottom of the taxonomy box to the right). Future taxonomic research is likely to conclude what some North American myrmecologists already believe - that this name subsumes more than the single species P. pilifera.

Identification
See the description in the nomenclature section.

Distribution
Pheidole pilifera is the most widespread and northward-reaching of all the Nearctic Pheidole. The species, as presently broadly construed, ranges from Massachusetts south to Georgia and west to California, extending through the midwest as far north as Minnesota, North Dakota and southwest to Utah, New Mexico, and Arizona. (Wilson 2003)

Distribution based on Regional Taxon Lists
Nearctic Region: United States. Neotropical Region: Mexico.

Biology
Colonies of pilifera excavate crater nests in exposed soil and harvest seeds, which are stored in nest chambers. In Colorado, Gregg (1963) found the species (as subsp. coloradensis) abundant at 1500 to 2600 m, in a wide range of habitats, from short-grass prairie, roadsides, and herbaceous semi-desert to mixed canyon forest and mountain mahogany shrub. Sexuals were present in the nest from the first week in June to the last week in July. P. pilifera is also notable in Colorado as the host of the social parasite. In Nevada, pilifera ranges between 900 and 2300 m, nesting in desert and juniper-pinyon woodland, both under stones and in the open soil, where it forms craters 25–60 mm wide (G. C. and J. Wheeler 1986). According to Stefan Cover (personal communication), “In the eastern U. S. pilifera occurs in open, grassy habitats, especially those with sandy soils containing a little clay. It is less common in pure sand, thus in pine barrens. Mating flights occur in early to mid July. Colonies are monogynous, and the newly mated queens start colonies singly. Minors are ‘shy’ foragers but recruit to good food sources. Majors seldom leave the nest except when recruited. Their primary function is to block nest passages, which they do effectively.” (Wilson 2003)

Worker
Minor

Nomenclature

 * pennsylvanica. Pheidole pennsylvanica Roger, 1863a: 199 (s) U.S.A. Mayr, 1886d: 456 (s.w.q.m.). Junior synonym of pilifera: Emery, 1895c: 290.
 *  pilifera. Leptothorax pilifer Roger, 1863a: 180 (w.) U.S.A. Emery, 1895c: 290 (s.); Wheeler, G.C. & Wheeler, J. 1953b: 79 (l.). Combination in Pheidole: Emery, 1895c: 290. Senior synonym of pennsylvanica: Emery, 1895c: 290; of septentrionalis and material of the unavailable name simulans referred here: Creighton, 1950a: 186; of artemisia, coloradensis, pacifica: Wilson, 2003: 589.
 * coloradensis. Pheidole pilifera var. coloradensis Emery, 1895c: 290 (s.w.) U.S.A. Wheeler, W.M. 1908e: 435 (q.m.). Subspecies of pilifera: Wheeler, W.M. 1908e: 435; Wheeler, G.C. & Wheeler, E.W. 1944: 244. Material of the unavailable name neomexicana referred here by Creighton, 1950a: 187. Junior synonym of pilifera: Wilson, 2003: 589.
 * septentrionalis. Pheidole pilifera subsp. septentrionalis Wheeler, W.M. 1908e: 437 (s.w.) U.S.A. Junior synonym of pilifera: Creighton, 1950a: 186.
 * pacifica. Pheidole xerophila subsp. pacifica Wheeler, W.M. 1915b: 404 (s.w.q.m.) U.S.A. Subspecies of pilifera: Creighton, 1950a: 187. Junior synonym of pilifera: Wilson, 2003: 589.
 * artemisia. Pheidole pilifera subsp. artemisia Cole, 1933: 616 (s.w.) U.S.A. Cole, 1938c: 372 (q.). Junior synonym of pilifera: Wilson, 2003: 589.

Description
From Wilson (2003): DIAGNOSIS A member of the “pilifera complex” of the larger pilifera group, comprising Pheidole calens, Pheidole californica, Pheidole carrolli, Pheidole cavigenis, Pheidole clementensis, Pheidole creightoni, Pheidole hoplitica, Pheidole littoralis, Pheidole micula, Pheidole pilifera, Pheidole polymorpha, Pheidole rugulosa, Pheidole senex, Pheidole soritis, Pheidole tepicana and Pheidole torosa, which complex is characterized by the following traits. Major: dorsal head surface extensively sculptured; occipital lobe horizontally rugulose (or, in carrolli smooth, in littoralis foveate, and in micula and soritis carinulate); postpetiole from above diamond-shaped, trapezoidal, or spinose. Minor: eye medium-sized to large. P. pilifera is distinguished within its complex by the following combination of traits.

Major: relatively large, HW about 1.6 mm; posterior dorsal profile of head straight or slightly concave; mesonotal convexity prominent and symmetrical in side view; propodeal spines robust, long, and nearly vertical to basal propodeal face; petiolar node in side view tapering apically to a blunt point; postpetiolar node from above very broad relative to petiolar node, and bluntly spinose.

Minor: eyes medium-sized; humerus in dorsal-oblique view subangulate; all of head posterior to the clypeus and mesosoma foveolate and opaque.

Closely related to Pheidole carrolli but very different in sculpturing of the major.

My synonymy of artemisia, coloradensis, and pacifica follows Creighton (1950a) in placing them as geographic variants, or subspecies, and thence here into synonymy at the species level. However, this assignment is not well documented and thus is regarded as provisional. The pattern of geographic variation within pilifera is as follows.

Western populations (“coloradensis”) ranging from North Dakota and Colorado to Nevada display a narrowing of the transverse rugulose band to the rearmost part of the major occiput and loss of the longitudinal carinulae on the vertex (anterior to the occiput), an area that is spotted with coarse foveae and shiny interspaces. The trend is climaxed in Nevada and California (“pacifica”) by replacement of the foveae on the vertex of the major by fine punctures. Southwestward, in Utah and Arizona (“artemisia”), the broad band of occipital rugulae of the major in eastern populations (“typical” pilifera) is retained, but the carinulae of the vertex are lost and replaced by a shiny surface.

MEASUREMENTS (mm) Major (Haddam, Connecticut): HW 1.60, HL 1.72, SL 0.74, EL 0.24, PW 0.68. Minor (Haddam): HW 0.54, HL 0.60, SL 0.52, EL 0.12, PW 0.36.

COLOR Major: light reddish brown.

Minor: medium reddish brown.



'''Figure. Upper: major. Lower: minor. CONNECTICUT: East Haddam. Scale bars = 1 mm.'''

Type Material
- as reported in Wilson (2003)

Etymology
Gr pilifera, hairy. (Wilson 2003)

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.
 * 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.
 * Annotated Ant Species List Ordway-Swisher Biological Station. Downloaded at http://ordway-swisher.ufl.edu/species/os-hymenoptera.htm on 5th Oct 2010.
 * Bestelmeyer B. T., and J. A. Wiens. 2001. Local and regional-scale responses of ant diversity to a semiarid biome transition. Ecography 24: 381-392.
 * Browne J. T., R. E. Gregg. 1969. A study of the ecological distribution of ants in Gregory Canyon, Boulder, Colorado. University of Colorado Studies. Series in Biology 30: 1-48
 * Buren W. F. 1944. A list of Iowa ants. Iowa State College Journal of Science 18:277-312
 * Campbell K. U., and T. O. Crist. 2017. Ant species assembly in constructed grasslands isstructured at patch and landscape levels. Insect Conservation and Diversity doi: 10.1111/icad.12215
 * Carroll T. M. 2011. The ants of Indiana (Hymenoptera: Formicidae). Master's Thesis Purdue university, 385 pages.
 * Cole A. C., Jr. 1933. Descriptions of two new ants of the genus Pheidole (Hymenoptera: Formicidae). Annals of the Entomological Society of America 26: 616-618.
 * Cole A. C., Jr. 1952. Notes on the Pheidole pilifera (Roger) complex and a description of a new subspecies (Hymenoptera: Formicidae). Journal of the Tennessee Academy of Science 27: 278-280.
 * Cole A. C., Jr. 1953. Studies of New Mexico ants. V. The genus Pheidole with synonymy (Hymenoptera: Formicidae). Journal of the Tennessee Academy of Science 28: 297-299.
 * Cole A. C., Jr. 1956. Observations of some members of the genus Pheidole in the southwestern United States with synonymy (Hymenoptera: Formicidae). Journal of the Tennessee Academy of Science 31: 112-118.
 * Coovert G. A. 2005. The Ants of Ohio (Hymenoptera: Formicidae). Ohio Biological Survey, Inc. 15(2): 1-207.
 * Coovert, G.A. 2005. The Ants of Ohio (Hymenoptera: Formicidae) Ohio Biological Survey Bulletin New Series Volume 15(2):1-196
 * 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
 * Dattilo W. et al. 2019. MEXICO ANTS: incidence and abundance along the Nearctic-Neotropical interface. Ecology https://doi.org/10.1002/ecy.2944
 * Davis W. T., and J. Bequaert. 1922. An annoted list of the ants of Staten Island and Long Island, N. Y. Bulletin of the Brooklyn Entomological Society 17(1): 1-25.
 * Del Toro I., K. Towle, D. N. Morrison, and S. L. Pelini. 2013. Community Structure, Ecological and Behavioral Traits of Ants (Hymenoptera: Formicidae) in Massachusetts Open and Forested Habitats. Northeastern Naturalist 20: 1-12.
 * Del Toro, I. 2010. PERSONAL COMMUNICATION. MUSEUM RECORDS COLLATED BY ISRAEL DEL TORO
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1052
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1053
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1054
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1055
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1056
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1057
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1058
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1059
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1061
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1062
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1063
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1064
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1065
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1066
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1067
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1068
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1069
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1070
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1071
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1072
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1073
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1074
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1075
 * DuBois M. B. 1985. Distribution of ants in Kansas: subfamilies Ponerinae, Ecitoninae, and Myrmicinae (Hymenoptera: Formicidae). Sociobiology 11: 153-1077
 * Dubois, M.B. and W.E. Laberge. 1988. An Annotated list of the ants of Illionois. pages 133-156 in Advances in Myrmecology, J. Trager
 * Emery C. 1895. Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna. (Schluss). Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 8: 257-360.
 * Frye J. A., T. Frye, and T. W. Suman. 2014. The ant fauna of inland sand dune communities in Worcester County, Maryland. Northeastern Naturalist, 21(3): 446-471.
 * General D., and L. Thompson. 2008. Ants of Arkansas Post National Memorial: How and Where Collected. Journal of the Arkansas Academy of Science 62: 52-60.
 * General D., and L. Thompson. 2008. New distributional records of ants in Arkansas. Journal of the Arkansas Academy of Science 62: 148-150.
 * General D.M. & Thompson L.C. 2008. New Distributional Records of Ants in Arkansas for 2008. Journal of the Arkansas Academy of Science. 63: 182-184
 * Graham, J.H., A.J. Krzysik, D.A. Kovacic, J.J. Duda, D.C. Freeman, J.M. Emlen, J.C. Zak, W.R. Long, M.P. Wallace, C. Chamberlin-Graham, J.P. Nutter and H.E. Balbach. 2008. Ant Community Composition across a Gradient of Disturbed Military Landscapes at Fort Benning, Georgia. Southeastern Naturalist 7(3):429-448
 * Gregg R. E. 1959. Key to the species of Pheidole (Hymenoptera: Formicidae) in the United States. Journal of the New York Entomological Society 66: 7-48.
 * Gregg, R.T. 1963. The Ants of Colorado.
 * Hill J. G. 2006. Field observations of Pheidole pilifera (Roger) and Paratrechina arenivaga (Wheeler) (Hymenoptera: Formicidae) in a Black Belt Prairie remnant in Mississippi. Journal of the Mississippi Academy of Sciences. 51: 183-186.
 * Hill J. V. 2012. The ant (Hymenoptera: Formicidae) fauna of the cedar glades and xeric limestone prairies of the Central Basin of Tennessee. Journal of the Tennessee Academy of Science 87(3): 135.
 * Hill J.G. & Brown R. L. 2010. The Ant (Hymenoptera: Formicidae) Fauna of Black Belt Prairie Remnants in Alabama and Mississippi. Southeastern Naturalist. 9: 73-84
 * Ivanov, K. 2019. The ants of Ohio (Hymenoptera, Formicidae): an updated checklist. Journal of Hymenoptera Research 70: 65–87.
 * 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. Personnal Database. Accessed on February 5th 2014 at http://www.asu.edu/clas/sirgtools/resources.htm
 * La Rivers I. 1968. A first listing of the ants of Nevada. Biological Society of Nevada, Occasional Papers 17: 1-12.
 * Lavigne R., and T. J. Tepedino. 1976. Checklist of the insects in Wyoming. I. Hymenoptera. Agric. Exp. Sta., Univ. Wyoming Res. J. 106: 24-26.
 * Lynch J. F. 1988. An annotated checklist and key to the species of ants (Hymenoptera: Formicidae) of the Chesapeake Bay region. The Maryland Naturalist 31: 61-106
 * MacGown, J. 2011. Ants of South Carolina (species list).
 * MacGown. J. 2011. Ants collected during the 25th Annual Cross Expedition at Tims Ford State Park, Franklin County, Tennessee
 * 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.P. and E. Mackay. XXXX. The Ants of New Mexico
 * Menke S. B., E. Gaulke, A. Hamel, and N. Vachter. 2015. The effects of restoration age and prescribed burns on grassland ant community structure. Environmental Entomology http://dx.doi.org/10.1093/ee/nvv110
 * 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.
 * 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.
 * Munsee, J. R.; Jansma, W. B.; Schrock, J. R. 1986. Revision of the checklist of Indiana ants with the addition of five new species (Hymenoptera: Formicidae). Proceedings of the Indiana Academy of Science 1985, publ. 1986 Vol. 95 pp. 265-274
 * Naves M. A. 1985. A monograph of the genus Pheidole in Florida, USA (Hymenoptera: Formicidae). Insecta Mundi 1: 53-90
 * Nuhn, T.P. and C.G. Wright. 1979. An Ecological Survey of Ants (Hymenoptera: Formicidae) in a Landscaped Suburban Habitat. American Midland Naturalist 102(2):353-362
 * 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.
 * 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.
 * Rivers, A., M. Barbercheck, B. Govaerts, and N. Verhulst. 2016. Conservation agriculture affects arthropod community composition in a rainfed maize-wheat system in central Mexico. Applied Soil Ecology 100: 81-90.
 * 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.
 * Roeder K. A., and D. V. Roeder. 2017. The Pheidole (Hymenoptera: Formicidae) of Oklahoma: new species records and distributional notes. Check List 13(2): 2071.
 * Smith M. R. 1924. An annotated list of the ants of Mississippi (Hym.). Entomological News 35: 47-54.
 * 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
 * Toennisson T. A., N. J. Sanders, W. E. Klingeman, and K. M. Vail. 2011. Influences on the Structure of Suburban Ant (Hymenoptera: Formicidae) Communities and the Abundance of Tapinoma sessile. Environ. Entomol. 40(6): 1397-1404.
 * Van Pelt A., and J. B. Gentry. 1985. The ants (Hymenoptera: Formicidae) of the Savannah River Plant, South Carolina. Dept. Energy, Savannah River Ecology Lab., Aiken, SC., Report SRO-NERP-14, 56 p.
 * Vásquez-Bolaños M. 2011. Lista de especies de hormigas (Hymenoptera: Formicidae) para México. Dugesiana 18: 95-133
 * Wheeler G. C., and E. W. Wheeler. 1944. Ants of North Dakota. North Dakota Historical Quarterly 11:231-271.
 * Wheeler G. C., and J. Wheeler J. 1989. A checklist of the ants of Oklahoma. Prairie Naturalist 21: 203-210.
 * Wheeler G. C., and J. Wheeler. 1986. The ants of Nevada. Los Angeles: Natural History Museum of Los Angeles County, vii + 138 pp.
 * Wheeler G. C., and J. Wheeler. 1987. A Checklist of the Ants of South Dakota. Prairie Nat. 19(3): 199-208.
 * Wheeler W. M. 1906. Fauna of New England. 7. List of the Formicidae. Occasional Papers of the Boston Society of Natural History 7: 1-24.
 * Wheeler W. M. 1908. The ants of Texas, New Mexico and Arizona. (Part I.). Bulletin of the American Museum of Natural History 24: 399-485.
 * Wheeler W. M. 1916. Formicoidea. Formicidae. Pp. 577-601 in: Viereck, H. L. 1916. Guide to the insects of Connecticut. Part III. The Hymenoptera, or wasp-like insects, of Connecticut. Connecticut State Geological and Natural History Survey. Bulletin 22: 1-824.
 * Wheeler, G.C. and J. Wheeler. 1985. A checklist of Texas ants. Prairie Naturalist 17:49-64.
 * Wheeler, G.C. and J. Wheeler. 1988. A checklist of the ants of Wyoming. Insecta Mundi 2(3&4):230-239
 * Wilson, E.O. 2003. Pheidole in the New World: A Dominant, Hyperdiverse Genus. Harvard University Press
 * 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