Abstract
This paper provides a taxonomic revision and a review of the biology for the 10 species of North American seed-harvester ants in the genus Veromessor. Two new synonomies are proposed: V. julianus subsp. clarior W.M. Wheeler and Creighton 1934 new synonym and V. julianus subsp. manni W.M. Wheeler and Creighton 1934 new synonym are synonomized under V. julianus (Pergande, 1894). One new species is described: V. pseudolariversi new species (worker, queen, male), as a result of splitting V. lariversi Smith into two species based on morphological and genetic differences. We also diagnose previously undescribed queens and males for the following species: V. andrei (male), V. chamberlini (queen, male), V. chicoensis (queen, male), V. julianus (queen, male), and V. stoddardi (queen, male). Information on biology of each species is summarized, along with distribution maps and keys to workers, queens, and males. We then discuss the biology for species of Veromessor, focusing on several morphological and ecological traits that display strong variation across the relatively low number of species (10) in the genus. Morphological traits include degree of psammophore development, propodeal spine length, eye size and structure, and worker polymorphism and worker body size, while ecological traits include colony size and foraging method, seasonality of mating flights, mating frequency, and queen size and colony founding strategy.
References
Andrews, H. (1916) A new ant of the genus Messor from Colorado. Psyche, 23 (3), 81–83.
Bennett, M. (2000) Systematics of the Nearctic granivore ant genus Veromessor. PhD dissertation, University of California, Davis, California, 114 pp.
Bernard, F. (1948) Les insectes sociaux du Fezzân. Comportement et biogéographie. In: Bernard, F. & de Peyerimhoff, P. (Eds.), Mission scientifique du Fezzân (1944–1945). Tome V. Zoologie. Institut de Recherches Sahariennes de l’Université d’Alger, Alger, pp. 87–200.
Blanchard, B.D. & Moreau, C.S. (2017) Defensive traits exhibit an evolutionary trade-off and drive diversification in ants. Evolution, 71 (2), 315–328. https://doi.org/10.1111/evo.13117
Blum, M.S., Padovani, F., Curley, A. & Hawk, R.E. (1969) Benzaldehyde: defensive secretion of a harvester ant. Comparative Biochemistry and Physiology, 29 (1), 461–465. https://doi.org/10.1016/0010-406X(69)91766-6
Bolton, B. (1982) Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax, Melissotarsus, Messor, and Cataulcus (Formicidae). Bulletin of the British Museum (Natural History) Entomology, 45 (4), 307–370.
Bolton, B. (1995) A New General Catalogue of the Ants of the World. Harvard University Press, Cambridge, Massachusetts, 504 pp.
Bolton, B. (2020) An online catalog of the ants of the world. Available from https://antcat.org, (accessed 18 September 2022)
Boudinot, B.E. (2013) The male genitalia of ants: musculature, homology, and functional morphology (Hymenoptera, Aculeata, Formicidae). Journal of Hymenoptera Research, 30, 29–49. https://doi.org/10.3897/jhr.30.3535
Boudinot, B.E., Moosdorf, O.T.D., Beutel, R.G. & Richter, A. (2021) Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): Patterns of sex- and caste-limited traits in the sausagefly and the driver ant. Journal of Morphology, 282 (11), 1616–1658. https://doi.org/10.1002/jmor.21410
Boulton, A.M., Jaffee, B.A. & Scow, K.M. (2003) Effects of a common harvester ant (Messor andrei) on richness and abundance of soil biota. Applied Soil Ecology, 23 (3), 257–265. https://doi.org/10.1016/S0929-1393(03)00046-5
Branstetter, M.G., Longino, J.T., Reyes-López,J.L., Brady, S.G. & Schultz, T.R. (2022). Out of the temperate zone: A phylogenomic test of the biogeographical conservatism hypothesis in a contrarian clade of ants. Journal of Biogeography, 49 (9):1640–1653. https://doi.org/10.1111/jbi.14462
Brown, M.J.F. (1999a) Nest relocation and encounters between colonies of the seed-harvesting ant Messor andrei. Insectes Sociaux, 46 (1), 66–70. https://doi.org/10.1007/s000400050114
Brown, M.J.F. (1999b) Semi-claustral founding and worker behaviour in gynes of Messor andrei. Insectes Sociaux, 46 (2), 194–195.
https://doi.org/10.1007/s000400050133
Brown, M.J.F. & Gordon, D.M. (2000) How resources and encounters affect the distribution of foraging activity in a seed-harvesting ant. Behavioral Ecology and Sociobiology, 47 (3), 195–203. https://doi.org/10.1007/s002650050011
Brown, M.J.F. & Human, K.G. (1997) Effects of harvester ants on plant species distribution and abundance in a serpentine grassland. Oecologia, 112 (2), 237–243. https://doi.org/10.1007/s004420050306
Brown, W.L. (1949) Synonomic and other notes on Formicidae. Psyche, 56 (1), 41–49.
Cahan, S. (2001a) Cooperation and conflict in ant foundress associations: insights from geographical variation. Animal Behaviour, 61 (4), 819–825. https://doi.org/10.1006/anbe.2000.1671
Cahan, S. (2001b) Ecological variation across a transition in colony-founding behavior in the ant Messor pergandei. Oecologia, 129 (4), 629–635. https://doi.org/10.1007/s004420100761
Cahan, S., Helms, K.R. & Rissing, S.W. (1998) An abrupt transition in colony founding behaviour in the ant Messor pergandei. Animal Behaviour, 55 (6), 1583–1594. https://doi.org/10.1006/anbe.1997.0699
Cahan, S. & Rissing, S.W. (2005) Variation in queen size across a behavioral transition zone in the ant Messor pergandei. Insectes Sociaux, 52 (1), 84–88. https://doi.org/10.1007/s00040-004-0776-7
Cantone, S. (2017) Winged Ants, the Male, Dichotomous Key to Genera of Winged Male Ants in the World, Behavioral Ecology of Mating Flight Autopubblicato, São Paulo, 318 pp. [https://www.academia.edu/35176438/Winged_Ants_The_Male_-Dichotomous_key_to_genera_of_winged_male_ants_in_the_World_Behavioral_Ecology_of_Mating_flight]
Clark, W.H. & Blom, P.E. (2007) Annotated checklist of the ants on the Idaho National Laboratory (Hymenoptera: Formicidae). Sociobiology, 49 (2), 1–117.
Cole, A.C. (1955) Studies of Nevada ants. I. Notes on Veromessor lariversi M.R. Smith and a description of the queen (Hymenoptera: Formicidae). Journal of the Tennessee Academy of Science, 30 (1), 51–52.
Cole, A.C. (1963) A new species of Veromessor from the Nevada test site and notes on related species (Hymenoptera: Formicidae). Annals of the Entomological Society of America, 56 (5), 678–682. https://doi.org/10.1093/aesa/56.5.678
Cole, A.C. (1966) Ants of the Nevada test site. Brigham Young University Science Bulletin - Biological Series, 7 (3), 1–27.
Creighton, W.S. (1950) The ants of North America. Bulletin of the Museum of Comparative Zoology, 104, 1–585.
Creighton, W.S. (1953) New data on the habits of the ants of the genus Veromessor. American Museum Novitates, 1612, 1–18.
Davidson, D.W. (1977) Species diversity and community organization in desert seed-eating ants. Ecology, 58 (4), 711–724. https://doi.org/10.2307/1936208
Davidson, D.W. (1978) Size variability in the worker caste of a social insect (Veromessor pergandei Mayr) as a function of the competitive environment. American Naturalist, 112 (985), 523–532. https://doi.org/10.1086/283294
Demarco, B.B. & Cognato, A.I. (2015) Phylogenetic analysis of Aphaenogaster supports the resurrection of Novomessor (Hymenoptera: Formicidae). Annals of the Entomological Society of America, 108 (2), 201–210. https://doi.org/10.1093/aesa/sau013
Denny, A.J., Franks, N.R., Powell, S. & Edwards, K.J. (2004) Exceptionally high levels of multiple mating in an army ant. Naturwissenschaften, 91 (8), 396–399. https://doi.org/10.1007/s00114-004-0546-4
do Nascimento, R.R., Jackson, B.D., Morgan, E.D., Clark, W.H. & Blom, P.E. (1993) Chemical secretions of two sympatric harvester ants, Pogonomyrmex salinus and Messor lobognathus. Journal of Chemical Ecology, 19 (9), 1993–2005. https://doi.org/10.1007/BF00983802
Donisthorpe, H. (1943) A list of the type-species of the genera and subgenera of the Formicidae. Annals and Magazine of Natural History, 11 (10), 721–737.
Emery, C. (1895) Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna. Zoologische Jahrbücher. Abtheilung fur Systematik, Geographie und Biologie der Tiere, 8, 257–360.
Emery, C. (1915) Definizione del genere Aphaenogaster e partizione di esso in sottogeneri. Parapheidole e Novomessor nn. gg. Rendiconti delle Sessioni della Reale Accademia delle Scienze dell’Istituto di Bologna. Classe di Scienze Fisiche, New Series, 19, 67–75.
Emery, C. (1921) Hymenoptera, Family Formicidae, Subfamily Myrmicinae [part]. Genera Insectorum, 174A, 1–94.
Enzmann, J. (1947) New forms of Aphaenogaster and Novomessor. Journal of the New York Entomological Society, 55 (2), 147–152.
Faircloth, B.C., McCormack, J.E., Crawford, N.G., Harvey, M.G., Brumfield, R.T. & Glenn, T.C. (2012) Ultraconserved elements anchor thousands of genetic markers spanning multiple evolutionary timescales. Systematic Biology, 61 (5), 717–726. https://doi.org/10.1093/sysbio/sys004
Feener, D.H., Jr. & Lighton, J.R.B. (1991) Is foraging in the desert ant, Messor pergandei (Hymenoptera: Formicidae), limited by water? Ecological Entomology, 16 (2), 183–191. https://doi.org/10.1111/j.1365-2311.1991.tb00208.x
Forel, A. (1917) Cadre synoptique actuel de la faune universelle des fourmis. Bulletin de la Société Vaudoise des Sciences Naturelles, 51, 229–253.
Freas, C.A., Congdon, J.V., Plowes, N.J.R. & Spetch, M.L. (2019) Same but different: socially foraging ants backtrack like individually foraging ants but use different mechanisms. Journal of Insect Physiology, 118, 103944. https://doi.org/10.1016/j.jinsphys.2019.103944
Freas, C.A., Congdon, J.V., Plowes, N.J.R. & Spetch, M.L. (2020) Pheromone cue triggers switch between vectors in the desert harvest ant, Veromessor pergandei. Animal Cognition, 23 (6), 1087–1105. https://doi.org/10.1007/s10071-020-01354-7
Freas, C.A., Plowes, N.J.R. & Spetch, M.L. (2019) Not just going with the flow: foraging ants attend to polarised light even while on the pheromone trail. Journal of Comparative Physiology A, 205 (5), 755–767. https://doi.org/10.1007/s00359-019-01363-z
Gordon, S.H. (1978) Food and foraging ecology of a desert harvester ant, Veromessor pergandei (Mayr). PhD dissertation, University of California, Berkeley, California, 186 pp.
Gregg, R.E. (1955) The rediscovery of Veromessor lobognathus (Andrews) (Hymenoptera: Formicidae). Psyche, 62 (2), 45–52. https://doi.org/10.1155/1955/50593
Greiner, B. (2006) Adaptations for nocturnal vision in insect apposition eyes. International Review of Cytology, 250, 1–46. https://doi.org/10.1016/S0074-7696(06)50001-4
Hagen, R.H., Smith, D.R. & Rissing, S.W. (1988) Genetic relatedness among co-foundresses of two desert ants, Veromessor pergandei and Acromyrmex versicolor (Hymenoptera: Formicidae). Psyche, 95 (3–4), 191–201.
Hahn, D.A., Johnson, R.A., Buck, N.A. & Wheeler, D.E. (2004) Storage protein content as a functional marker for colony-founding strategies: a comparative study within the harvester ant genus, Pogonomyrmex. Physiological and Biochemical Zoology, 77 (1), 100–108. https://doi.org/10.1086/380214
Hale, A., Bougie, T., Henderson, E., Sankovitz, M., West, M. & Purcell, J. (2018) Notes on hunting behavior of the spider Euryopis californica Banks, 1904 (Araneae: Theridiidae), a novel predator of Veromessor pergandei (Mayr, 1886) harvester ants (Hymenoptera: Formicidae). Pan-Pacific Entomologist, 93 (3), 141–145. https://doi.org/10.3956/2018-94.3.141
Hamm, C.A. & Kamansky, B. (2009) New record of Messor chicoensis (Hymenoptera: Formicidae) from the San Joaquin Valley of California. Sociobiology, 53 (2), 543–547.
Helms Cahan, S. & Helms, K.R. (2012) Relatedness does not explain geographic variation in queen cooperation in the seed-harvester ant Messor pergandei. Insectes Sociaux, 59 (4), 579–585. https://doi.org/10.1007/s00040-012-0255-5
Helms Cahan, S. & Helms, K.R. (2014) Variation in social structure alters queen body mass and productivity in the desert seed-harvester ant Messor pergandei. Insectes Sociaux, 61 (2), 153–161. https://doi.org/10.1007/s00040-014-0340-z
Helms, K.R. & Helms Cahan, S. (2012) Large-scale regional variation in cooperation and conflict among queens of the desert ant Messor pergandei. Animal Behaviour, 84 (3), 499–507. https://doi.org/10.1016/j.anbehav.2012.05.019
Helms, K.R., Newman, N.J. & Helms Cahan, S. (2013) Regional variation in queen and worker aggression in incipient colonies of the desert ant Messor pergandei. Behavioral Ecology and Sociobiology, 67 (10), 1563–1573. https://doi.org/10.1007/s00265-013-1568-8
Hobbs, R.J. (1985) Harvester ant foraging and plant species distribution. Oecologia, 67 (4), 519–523. https://doi.org/10.1007/BF00790023
Hölldobler, B., Plowes, N.J.R., Johnson, R.A., Nishshanka, U., Liu, C. & Attygalle, A.B. (2013) Pygidial gland chemistry and potential alarm-recruitment function in column foraging, but not solitary, Nearctic Messor harvesting ants (Hymenoptera: Formicidae: Myrmicinae). Journal of Insect Physiology, 59 (9), 863–869. https://doi.org/10.1016/j.jinsphys.2013.06.006
Johnson, R.A. (1989) The ecology and evolution of learning in the seed-harvester ants Veromessor pergandei and Pogonomyrmex rugosus. PhD dissertation, Arizona State University, Tempe, Arizona, 138 pp.
Johnson, R.A. (1991) Learning, memory, and foraging efficiency in two species of desert seed-harvester ants. Ecology, 72 (4), 1408–1419. https://doi.org/10.2307/1941112
Johnson, R.A. (1992) Soil texture as an influence on the distribution of the desert seed-harvester ants Pogonomyrmex rugosus and Messor pergandei. Oecologia, 89 (1), 118–124. https://doi.org/10.1007/BF00319023
Johnson, R.A. (1998) Foundress survival and brood production in the desert seed-harvester ants Pogonomyrmex rugosus and P. barbatus (Hymenoptera, Formicidae). Insectes Sociaux, 45 (3), 255–266. https://doi.org/10.1007/s000400050086
Johnson, R.A. (2000a) 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. (2000b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89–122.
Johnson, R.A. (2000c) Water loss in desert ants: caste variation and the effect of cuticle abrasion. Physiological Entomology, 25 (1), 48–53. https://doi.org/10.1046/j.1365-3032.2000.00170.x
Johnson, R.A. (2001) Biogeography and community structure of North American seed-harvester ants. Annual Review of Entomology, 46, 1–29. https://doi.org/10.1146/annurev.ento.46.1.1
Johnson, R.A. (2002) Semi-claustral colony founding in the seed-harvester ant Pogonomyrmex californicus: a comparative analysis of founding strategies. Oecologia, 132 (1), 60–67. https://doi.org/10.1007/s00442-002-0960-2
Johnson, R.A. (2006) Capital and income breeding and the evolution of colony founding strategies in ants. Insectes Sociaux, 53 (3), 316–322. https://doi.org/10.1007/s00040-006-0874-9
Johnson, R.A. (2015) A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part I. Zootaxa, 4029 (1), 1–142. https://doi.org/10.11646/zootaxa.4029.1.1
Johnson, R.A. (2021) Dessication limits recruitment in the pleometrotic desert seed-harvester ant Veromessor pergandei. Ecology and Evolution, 11 (1), 294–308. https://doi.org/10.1002/ece3.7039
Johnson, R.A. & Cover, S.P. (2015) A taxonomic revision of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae) on Hispaniola. Zootaxa, 3972 (2), 231–249. https://doi.org/10.11646/zootaxa.3972.2.5
Johnson, R.A., Holbrook, C.T., Strehl, C. & Gadau, J. (2007) Population and colony structure and morphometrics in the queen dimorphic harvester ant, Pogonomyrmex pima. Insectes Sociaux, 54 (1), 77–86. https://doi.org/10.1007/s00040-007-0916-y
Johnson, R.A., Kaiser, A., Quinlan, M. & Sharp, W. (2011) Effect of cuticular abrasion and recovery on water loss rates in queens of the desert harvester ant Messor pergandei. Journal of Experimental Biology, 214 (20), 3495–3506. https://doi.org/10.1242/jeb.054304
Johnson, R.A., Rissing, S.W. & Killeen, P.R. (1994) Differential learning and memory by co-occurring ant species. Insectes Sociaux, 41 (2), 165–177. https://doi.org/10.1007/BF01240476
Johnson, R.A. & Rutowski, R.L. (2022) Color, activity period, and eye structure in four lineages of ants: pale, nocturnal species have evolved larger eyes and larger facets than their dark, diurnal congeners. PLoS ONE, 17 (9), e0257779. https://doi.org/10.1371/journal.pone.0257779
Johnson, R.A. & Ward, P.S. (2002) Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: a first overview. Journal of Biogeography, 29 (8), 1009–1026. https://doi.org/10.1046/j.1365-2699.2002.00746.x
Jones, G. (2017) Algorithmic improvements to species delimitation and phylogeny estimation under the multispecies coalescent. Journal of Mathematical Biology, 74 (1–2), 447–467. https://doi.org/10.1007/s00285-016-1034-0
Keller, R.A. (2011) A phylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the Poneromorph subfamilies. Bulletin of the American Museum of Natural History, 355, 1–90. https://doi.org/10.1206/355.1
Krebs, R.A. & Rissing, S.W. (1991) Preference for larger foundress associations in the desert ant Messor pergandei. Animal Behaviour, 41 (2), 361–363. https://doi.org/10.1016/S0003-3472(05)80487-7
Kwapich, C.L., Gadau, J. & Hölldobler, B. (2017) The ecological and genetic basis of annual worker production in the desert seed harvesting ant, Veromessor pergandei. Behavioral Ecology and Sociobiology, 71, Article 110. https://doi.org/10.1007/s00265-017-2333-1
Kwapich, C.L. & Hölldobler, B. (2019) Destruction of spider webs and rescue of ensnared nestmates by a granivorous desert ant (Veromessor pergandei). American Naturalist, 194 (3), 395–404. https://doi.org/10.1086/704338
Kwapich, C.L., Valentini, G. & Hölldobler, B. (2018) The non-additive effects of body size on nest architecture in a polymorphic ant. Philosophical Transactions of the Royal Society, Series B, 373, 20170235. https://doi.org/10.1098/rstb.2017.0235
Lighton, J.R.B. & Berrigan, D. (1995) Questioning paradigms: caste-specific ventilation in harvester ants, Messor pergandei and M. julianus (Hymenoptera: Formicidae). Journal of Experimental Biology, 198 (2), 521–530. https://doi.org/10.1242/jeb.198.2.521
Lighton, J.R.B., Quinlan, M.C. & Feener, D.H., Jr. (1994) Is bigger better? Water balance in the polymorphic desert harvester ant Messor pergandei. Physiological Entomology, 19 (4), 325–334. https://doi.org/10.1111/j.1365-3032.1994.tb01059.x
Mayr, G. (1886) Die Formiciden der Vereinigten Staaten von Nordamerika. Verhandlungen Zoologisch-Botanische Gesellschaft in Wein, 36, 419–464.
McCluskey, E.S. (1958) Daily rhythms in male harvester and Argentine ants. Science, 128 (3323), 536–537. https://doi.org/10.1126/science.128.3323.536
McCluskey, E.S. (1963) Rhythms and clocks in harvester and Argentine ants. Physiological Zoology, 36 (3), 273–292.
Muscedere, M.L., Helms Cahan, S., Helms, K.R. & Traniello, J.F.A. (2016) Geographic and life-history variation in ant queen colony founding correlates with brain amine levels. Behavioral Ecology, 27 (1), 271–278. https://doi.org/10.1093/beheco/arv152
Ode, P.J. & Rissing, S.W. (2002) Resource abundance and sex allocation by queen and workers in the harvester ant, Messor pergandei. Behavioral Ecology and Sociobiology, 51 (6), 548–556. https://doi.org/10.1007/s00265-002-0462-6
Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’Amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: a new map of life on Earth. Bioscience, 51 (11), 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
Overson, R.P., Fewell, J.H. & Gadau, J. (2016) Distribution and origin of intraspecific social variation in the California harvester ant Pogonomyrmex californicus. Insectes Sociaux, 63 (4), 531–541. https://doi.org/10.1007/s00040-016-0497-8
Peeters, C. (1997) Morphologically ‘primitive’ ants: comparative review of social characters, and the importance of queen-worker dimorphism. In: Choe, J.C. & Crespi, B.J. (Eds.), The Evolution of Social Behavior in Insects and Arachnids. Cambridge University Press, Cambridge, pp. 372–391. https://doi.org/10.1017/CBO9780511721953.019
Pergande, T. (1894) Formicidae of Lower California, Mexico. Proceedings of the California Academy of Sciences, 4 (2), 161–165.
Peters, H.A., Chiariello, N.R., Mooney, H.A., Levin, S.A. & Hartley, A.E. (2005) Native harvester ants threatened with widespread displacement exert localized effects on serpentine grassland plant community composition. Oikos, 109 (2), 351–359. https://doi.org/10.1111/j.0030-1299.2005.13282.x
Pfennig, D.W. (1995) Absence of joint nesting advantage in desert seed harvester ants: evidence from a field experiment. Animal Behaviour, 49 (3), 567–575. https://doi.org/10.1016/0003-3472(95)80190-1
Pinter-Wollman, N. & Brown, M.J.F. (2015) Variation in nest relocation of harvester ants is affected by population density and food abundance. Behavioral Ecology, 26 (6), 1569–1576. https://doi.org/10.1093/beheco/arv108
Plowes, N., Du., Y., Congdon, J.V., Bulitko, V., Soares, E.S. & Spetch, M.L. (2019) Odometry and backtracking: social and individual navigation in group foraging desert harvester ants (Veromessor pergandei). Animal Cognition, 22 (1), 35–47. https://doi.org/10.1007/s10071-018-1218-7
Plowes, N.J.R., Colella, T., Johnson, R.A. & Hölldobler, B. (2014) Chemical communication during foraging in the harvesting ants Messor pergandei and Messor andrei. Journal of Comparative Physiology A, 200 (2), 129–137. https://doi.org/10.1007/s00359-013-0868-9
Plowes, N.J.R., Johnson, R.A. & Hölldobler, B. (2012) Foraging behavior in the ant genus Messor (Hymenoptera: Formicidae: Myrmicinae). Myrmecological News, 18, 33–49.
Plowes, N.J.R., Ramsch, K., Middendorf, M. & Hölldobler, B. (2014) An empirically based simulation of group foraging in the harvesting ant, Messor pergandei. Journal of Theoretical Biology, 340, 186–198. https://doi.org/10.1016/j.jtbi.2013.07.014
Pollock, G.B. & Rissing, S.W. (1985) Mating season and colony foundation of the seed-harvester ant, Veromessor pergandei. Psyche, 92 (1), 125–134. https://doi.org/10.1155/1985/87410
Porter, S.D. & Jorgensen, C.D. (1990) Psammophores: do harvester ants (Hymenoptera: Formicidae) use these pouches to transport seeds? Journal of the Kansas Entomological Society, 63 (1), 138–149.
Raczkowski, J.M. (2003) Advantage of pleometrosis in Messor pergandei (Formicidae: Myrmicinae): evidence from field experiments. MS thesis, Ohio State University, Columbus, Ohio, 42 pp.
Rissing, S.W. (1981) Foraging specializations of individual seed-harvester ants. Behavioral Ecology and Sociobiology, 9 (2), 149–152. https://doi.org/10.1007/BF00293586
Rissing, S.W. (1986) Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests. Oecologia, 68 (2), 231–234. https://doi.org/10.1007/BF00384792
Rissing, S.W. (1987) Annual cycles in worker size of the seed-harvester ant Veromessor pergandei (Hymenoptera: Formicidae). Behavioral Ecology and Sociobiology, 20 (2), 117–124. https://doi.org/10.1007/BF00572633
Rissing, S.W. (1988) Dietary similarity and foraging range of two seed-harvester ants during resource fluctuations. Oecologia, 75 (3), 362–366. https://doi.org/10.1007/BF00376938
Rissing, S.W., Johnson, R.A. & Martin, J.W. (2000) Colony founding behavior of some desert ants: geographic variation in metrosis. Psyche, 103 (1–2), 95–101.
Rissing, S.W., Johnson, R.A. & Pollock, G.B. (1986) Natal nest distribution and pleometrosis in the desert leaf-cutter ant Acromyrmex versicolor (Hymenoptera: Formicidae). Psyche, 93 (3–4), 177–186.
Rissing, S.W. & Pollock, G.B. (1984) Worker size variability and foraging efficiency in Veromessor pergandei (Hymenoptera: Formicidae). Behavioral Ecology and Sociobiology, 15 (2), 121–126. https://doi.org/10.1007/BF00299379
Rissing, S.W. & Pollock, G.B. (1986) Social interaction among pleometrotic queens of Veromessor pergandei (Hymenoptera: Formicidae) during colony foundation. Animal Behaviour, 34 (1), 226–233. https://doi.org/10.1016/0003-3472(86)90027-8
Rissing, S.W. & Pollock, G.B. (1987) Queen aggression, pleometrotic advantage and brood raiding in the ant Veromessor pergandei (Hymenoptera: Formicidae). Animal Behaviour, 35 (4), 975–981. https://doi.org/10.1016/S0003-3472(87)80154-9
Rissing, S.W. & Pollock, G.B. (1989) Behavioral ecology and community organization of desert seed-harvester ants. Journal of Arid Environments, 17 (2), 167–173. https://doi.org/10.1016/S0140-1963(18)30903-0
Rissing, S.W. & Pollock, G.B. (1991) An experimental analysis of pleometrotic advantage in the desert seed-harvester ant Messor pergandei (Hymenoptera; Formicidae). Insectes Sociaux, 38 (2), 205–211. https://doi.org/10.1007/BF01240970
Rissing, S.W. & Wheeler, J. (1976) Foraging responses of Veromessor pergandei to changes in seed production. Pan-Pacific Entomologist, 52 (1), 63–72.
Ryti, R.T. (1988) Geographic variation in cooperative colony foundation in Veromessor pergandei (Hymenoptera: Formicidae). Pan-Pacific Entomologist, 64 (3), 255–257.
Ryti, R.T. & Case, T.J. (1984) Spatial arrangement and diet overlap between colonies of desert ants. Oecologia, 62 (3), 401–404. https://doi.org/10.1007/BF00384274
Ryti, R.T. & Case, T.J. (1986) Overdispersion of ant colonies: a test of hypotheses. Oecologia, 69 (3), 446–453. https://doi.org/10.1007/BF00377067
Ryti, R.T. & Case, T.J. (1988a) Field experiments on desert ants: testing for competition between colonies. Ecology, 69 (6), 1993–2003. https://doi.org/10.2307/1941177
Ryti, R.T. & Case, T.J. (1988b) The regeneration niche of desert ants: effects of established colonies. Oecologia, 75 (2), 303–306.
https://doi.org/10.1007/BF00378614
Sarnat, E.M., Friedman, N.R., Fischer, G., Lecroq-Bennet, B. & Economo, E.P. (2017) Rise of the spiny ants: diversification, ecology and function of extreme traits in the hyperdiverse genus Pheidole (Hymenoptera: Formicidae). Biological Journal of the Linnean Society, 122 (3), 514–538. https://doi.org/10.1093/biolinnean/blx081
Smith, M.R. (1951) Two new ants from western Nevada. Great Basin Naturalist, 11 (3–4), 91–96.
Smith, M.R. (1956) A key to the workers of Veromessor Forel of the United States and the description of a new subspecies. Pan-Pacific Entomologist, 32 (1), 36–38.
Snelling, R.R. & George, C.D. (1979) The taxonomy, distribution and ecology of California desert ants (Hymenoptera: Formicidae). Report to Bureau of Land Management, US Department of Interior, Riverside, California, 335 + 89 pp.
Spangler, H.G. & Rettenmeyer, C.W. (1966) The function of the ammochaetae or psammophores of harvester ants, Pogonomyrmex spp. Journal of the Kansas Entomological Society, 39 (4), 739–745.
Stöckl, A., Smolka, J., O’Carroll, D. & Warrant, E. (2017) Resolving the trade-off between visual sensitivity and spatial acuity–lessons from hawkmoths. Integrative and Comparative Biology, 57 (5), 1093–1103. https://doi.org/10.1093/icb/icx058
Sukumaran, J., Holder, M.T. & Knowles, L.L. (2021) Incorporating the speciation process into species delimitation. PLoS Computational Biology, 17 (15), e1008924. https://doi.org/10.1371/journal.pcbi.1008924
Taber, S.W. & Cokendolpher, J.C. (1988) Karyotypes of a dozen ant species from the southwestern U.S.A. (Hymenoptera: Formicidae). Caryologia, 41 (2), 93–102. https://doi.org/10.1080/00087114.1988.10797851
Tevis, L. Jr. (1958) Interrelations between the harvester ant Veromessor pergandei (Mayr) and some desert ephemerals. Ecology, 39 (4), 695–704. https://doi.org/10.2307/1931610
Tschinkel, W.R. (2015) The architecture of subterranean nests: beauty and mystery underfoot. Journal of Bioeconomics, 17 (3), 271–291. https://doi.org/10.1007/s10818-015-9203-6
Villesen, P., Murakami, T., Schultz, T.R. & Boomsma, J.J. (2002) Identifying the transition between single and multiple mating of queens in fungus-growing ants. Proceeding of the Royal Society of London, Series B, 269 (1500), 1541–1548. https://doi.org/10.1098/rspb.2002.2044
Ward, P.S., Brady, S.G., Fisher, B.L. & Schultz, T.R. (2015) The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology, 40 (1), 61–81. https://doi.org/10.1111/syen.12090
Waser, N.M. (1998) Task-matching and short-term size shifts in foragers of the harvester ant, Messor pergandei (Hymenoptera: Formicidae). Journal of Insect Behavior, 11 (3), 451–462. https://doi.org/10.1023/A:1020963000112
Went, F.W., Wheeler, J. & Wheeler, G.C. (1972) Feeding and digestion in some ants (Veromessor and Manica). Bioscience, 22 (2), 82–88. https://doi.org/10.2307/1296037
Wheeler, G.C. & Wheeler, J. (1956) Veromessor lobognathus in North Dakota (Hymenoptera: Formicidae). Psyche, 63 (4), 140–145.
Wheeler, G.C. & Wheeler, J. (1959) Veromessor lobognathus: second note (Hymenoptera: Formicidae). Annals of the Entomological Society of America, 52 (2), 176–179.
Wheeler, G.C. & Wheeler, J. (1965) Veromessor lobognathus: third note (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society, 38 (1), 55–61.
Wheeler, G.C. & Wheeler, J. (1967) Veromessor lobognathus: fourth note (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society, 40 (2), 238–241.
Wheeler, G.C. & Wheeler, J. (1972) Ant larvae of the subfamily Myrmicinae: second supplement on the tribes Myrmicini and Pheidolini. Journal of the Georgia Entomological Society, 7, 233–246.
Wheeler, G.C. & Wheeler, J. (1973) Ants of Deep Canyon. University of California Press, Berkeley, California, 162 pp.
Wheeler, G.C. & Wheeler, J.N. (1986) The Ants of Nevada. Natural History Museum of Los Angeles County, Los Angeles, California, 138 pp.
Wheeler, J. & Rissing, S.W. (1975a) Natural history of Veromessor pergandei. I. The nest. Pan-Pacific Entomologist, 51 (3), 205–216.
Wheeler, J. & Rissing, S.W. (1975b) Natural history of Veromessor pergandei. II. Behavior. Pan-Pacific Entomologist, 51 (4), 303–314.
Wheeler, W.M. (1907) On certain modified hairs peculiar to the ants of arid regions. Biological Bulletin, 13 (4), 185–202. https://doi.org/10.2307/1535694
Wheeler, W.M. (1915) Some additions to the North American ant-fauna. Bulletin of the American Museum of Natural History, 34, 389–421.
Wheeler, W.M. (1922) Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631–710.
Wheeler, W.M. & Creighton, W.S. (1934) A study of the ant genera Novomessor and Veromessor. Proceedings of the American Academy of Arts and Sciences, 69 (9), 341–387. https://doi.org/10.2307/20023057
Wiernasz, D.C. & Cole, B.J. (2003) Queen size mediates queen survival and colony fitness in harvester ants. Evolution, 57 (9), 2179–2183. https://doi.org/10.1111/j.0014-3820.2003.tb00396.x
Wiernasz, D.C., Perroni, C.L. & Cole, B.J. (2004) Polyandry and fitness in the western harvester ant, Pogonomyrmex occidentalis. Molecular Ecology, 13 (6), 1601–1606. https://doi.org/10.1111/j.1365-294X.2004.02153.x
Wills, B.D., Powell, S., Rivera, M.D. & Suarez, A.V. (2018) Correlates and consequences of polymorphism in ants. Annual Review of Entomology, 63, 575–598. https://doi.org/10.1146/annurev-ento-020117-043357
Wilson, E.O. (1953) The origin and evolution of polymorphism in ants. Quarterly Review of Biology, 28 (2), 136–156. https://doi.org/10.1086/399512