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Superfamily FORMICOIDEA
Family FORMICIDAE

Ants are social insects that live in colonies in various ecological situations, most commonly in
the soil, rotting wood, and plant cavities. They are practically ubiquitous being extremely prolific
in numbers of individuals even though relatively low in numbers of species. Although most ants
are free-living, some are parasites on other sjjecies of ants or live as inquilines in the nests of
other ants. Most ants are omnivorous, but many have a more highly specialized food require-
ment. A number of forms are of concern to man because of their feeding, nesting, or stinging
habits. The species that nest in or near houses or other buildings are sometimes considerably an-
noying to man. Because of the adaptive ability of many species and their habits of nesting in
plants and soil, many forms have been spread throughout the world by commerce. A number of
these tramp species are found in North America.

A colony of ants is usually composed of one or more reproductive females (queens) and wor-
kers. At certain times of the year a nest also contains males and virgin females. There are three
distinct castes of ants, the female, males, and workers. The female is winged but loses her wings
soon after finding a suitable place to begin a new colony; her main purpose is reproduction. The
male is winged and is short-lived; he dies soon after mating. The workers are not winged, and
their main functions are to build the nest, feed the colony, care for the young, and defend the
nest. In some cases, workers are also reproductive forms. Many ants are monomorphic, that is,
the workers are all the same size. Other ants are polymorphic, in which case the workers vary
considerably in size. In polymorphic sjjecies, the largest workers are commonly termed soldiers
or maxims and the smaller workers termed minors or minims. The size of a colony of ants, de-
pending on the species and age of the colony, varies and may contain only a few dozen to many
thousands of individuals.

The literature on ants in voluminous. It is impossible to cite every reference on the subject.
The following list contains some of the more significant works on a variety of subjects. Further
references may be found in the literature cited sections of these citations. There have been
many local or state faunal lists and treatments of ants; many of these are given by Smith (1947)
who listed 84 publications by states. They are not repeated here.

I appreciate the cooperation of the following myrmecologists who have reviewed all or parts
of this section: A. C. Cole, University of Tennessee, Knoxville; A. Francoeur, Universite du
Quebec a Chicoutimi; M. R. Smith, Arlington, Virginia; R. R. Snelling, Los Angeles County
Museum, California; and J. F. Watkins II, Baylor University, Waco, Texas. This section is based
in large part on the Formicidae sections of the first Hymenoptera Catalog (1951) and its supple-
ments (1958, 1967) by M. R. Smith.

Revision: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., BuL 104: 1-585, 57 pis. (keys to
genera and species of N. Amer.).




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Taxonomy: Dalla Torre, 1893. Cat. Hym., v. 7, 289 pp. (world cat.). —Emery, 1910. In
Wytsman, Gen. Ins., fasc. 102, 34 pp. (Dorylinae, world cat.). — Emery, 1911. In Wytsman,
Gen. Ins., fasc. 118, 124 pp. (Ponerinae, world cat.). — Wheeler, 1911. N. Y. Acad. Sci., Ann.
21: 157-175 (type-species). —Emery, 1912. In Wytsman, Gen. Ins., fasc. 137, 50 pp.
(Dolichoderinae, world cat.). — Wheeler, 1920. Psyche 27: 46-55 (subfamilies). — Emery,
1921. In Wytsman, Gen. Ins., fasc. 174, 379 pp. (Myrmicinae, world cat.). —Wheeler, 1922.
Amer. Mus. Nat. Hist., Bui. 45: 631-710 (keys to world genera and subgenera). — Emery,
1925. In Wytsman, Gen. Ins., fasc. 183, 302 pp. (Formicinae, world cat.). —Carpenter, 1930.
Harvard Univ., Mus. Comp. Zool., Bui. 70: 1-66 (fossil ants of N. Amer.). — Donisthorpe,
1943. Ann. and Mag. Nat. Hist. (11) 10: 617-648, 649-688, 721-737 (type-species). —Smith,
1943. Amer. Midland Nat. 30: 273-321 (key to N. Amer. genera based on males). —Smith,
1947. Amer. Midland Nat. 37: 521-647 (key to N. Amer. genera based on workers; list of
publications on ants by states). —Van Pelt, 1948. Fla. Ent. 30: 57-67 (spp. of Alachua Co.,
Fla.). — Brown, 1954. Insectes Sociaux 1: 21-31 (phylogeny and subfamily classification).
—Van Pelt, 1956. Amer. Midland Nat. 56: 358-387 (spp. of Welaka Reserve, Fla.).
— Kannowski, 1956. Amer. Midland Nat. 56: 168-185 (spp. of Ramsey Co., N. Dak.). —Van
Pelt, 1958. Amer. Midland Nat. 59: 1-57 (list of spp. of Welaka Reserve, Fla.). — Wenner,
1959. Amer. Midland Nat. 62: 174-182 (spp. of Bidwell Park, Chico, Calif.). —Wheeler and
Wheeler, 1960. Psyche 67: 87-94 (techniques for study of larvae). —Carter, 1962. Elisha
Mitchell Sci. Soc., Jour. 78: 150-204 (N. C). —Carter, 1962. EUsha MitcheU Sci. See., Jour.
78: 1-18 (N. C. Piedmont). —Wheeler and Wheeler, 1963. Ants of N. Dak, Univ. N. Dak.
Press, Grand Forks, 326 pp. —Gregg, 1963. Ants of Colo., Univ. Colo. Press, Boulder, 792
pp. — WUson, 1964. Breviora 210: 1-14 (ants of Fla. Keys). —Young and Howell, 1964. Okla.
Agr. Expt. Sta. MP 71 (spp. of Okla.). —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol.
Ser. 7 (3): 1-27 (spp. of Nev. Test Site). —WUson and Taylor, 1967. Pacific Ins. Monog. 14:
1-109 (spp. of Polynesia). —Wilson, Carpenter, and Brown, 1967. Psyche 74: 1-19 (first
Mesozoic ant; a new subfamily). —Warren and Rouse, 1969. Ark. Univ., Agr. Expt. Sta.,
Div. Agr. Bui. 742, 67 pp. (spp. of Ark). —Ross, Rotramel, and LaBerge, 1971. 111. Nat.
Hist. Survey, Biol. Notes No. 71, 22 pp. (common and economic ants of 111.). — Wheeler and
Wheeler, 1972. Ent. Soc. Wash., Proc. 74: 35-45 (subfamiUes). — Kempf, 1972. Studia Ent.
15: 3-344 (cat. of Neotropical ants). — Wheeler and Wheeler, 1973. Ants of Deep Canyon,
Philip L. Boyd Deep Canyon Research Center, Univ. Calif., Riverside, 159 pp. — Brown,
1973. In Meggers, et al.. Tropical forest ecosystems in Africa and S. Amer., Smithsn. Inst.
Press, Wash., D. C, pp. 161-185 (list of world generic and subgeneric names; suggested
synonymy). —Wheeler and Wheeler, 1976. Ent. Soc. Wash., Mem. 7, 108 pp. (ant larvae:
review and synthesis).

Biology: McCook, 1882. The honey ants of the Garden of the (Jods, and the Occident ants of
the American plains, Philadelphia, 188 pp. — Buckingham, 1911. Amer. Acad. Arts and Sci.,
Proc. 46: 425-507 (division of labor). — Wheeler, 1910. Ants, their structure, development,
and behavior, Columbia Univ. Press, N. Y., 663 pp. (reprinted 1926, 1960). —Wheeler, 1928.
The social insects, their origin and evolution, Harcourt, Brace and Co., N. Y., 378 pp.
— Forel, 1928. The social world of the ants, G. P. Putnam's Sons, Ltd., London and N. Y.,
V. I, 551 pp., v. II, 445 pp. —Jones, 1929. Colo. Agr. Col., Expt. Sta. Bui. 341, 96 pp.
(relation to aphids). —Wheeler, 1936. Amer. Acad. Arts and Sci., Proc. 71: 159-243
(ecological relations to termites). — Haskins and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37:
97-162 (sociological an physiological features). — Haskins, 1939. Of ants and men,
Prentice-Hall Inc., N. Y., 244 pp. — Lafleur, 1941. N. Y. Ent. Soc., Jour. 49: 227-231 (civil
disturbances in ant communities). — Lafleur, 1941. N. Y. Ent. Soc., Jour. 49: 199-204
(communal disaffection). — Lafleur, 1942. Sci. Monthly 65: 467^71 (behavior in the face of
obstacles). —Smith and Weiss, 1942. U. S. Dept. Agr., Tech. Bui. 798, 44 pp. (relation to
azalea flower spot). — Wheeler, 1942. Harvard Univ., Mus. Comp. Zool., Bui. 40: 1-252
(Neotropical ant-plants and their ants). — MacGregor, 1948. Behavioiu* 1: 267-296 (odor as
a basis for orientated movement). — Brackbill, 1948. Auk 65: 66-77 (anting by birds).
— Nixon, 1951. The association of ants with aphids and coccids, Commonwealth Inst. Ent.
(London). 35 pp. — Valentini, 1951. Ann. Sci. Nat. (Zool.) 11: 249-276 (adaptation of larvae).
— Flanders, 1951. Canad. Ent. 83: 93-98 (role of ants in biological control of homopterous
insects). — Flanders, 1952. Jour. Econ. Ent. 45: 38-39 (ovisorption as the mechanism




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causing worker development). — Flanders, 1953. Sci. Monthly 76: 142-148 (caste
determination). —Talbot, 1953. Mich. Univ., Contrib. Lab. Vertebrate Biol. 63, 13 pp.
(fauna and populations). — Wilson, 1953. Psyche 60: 15-20 (caste determination). — Simeone,
1954. State Univ. N. Y., Syracuse, Col. Forestry Bui. 34, 19 pp. (carpenter ants and their
control). — Chapman, 1954. Pan-Pacific Ent. 30: 93-102 (swarming of ants on mountain
summits). — Gosswald, 1955. Rev. Suisse de Zool. 62: 372-386 (caste determination).
— O'Rourke, 1956. Insectes Sociaux 3: 107-118 (medical and veterinary importance).
— Wilson and Eisner, 1957. Insectes Sociaux 4: 157-166 (liquid transmission of food).
—Talbot, 1957. Insectes Sociaux 4: 375-384 (populations). —Brian, 1957. Ann.Rev. Ent. 2:
107-120 (caste determination). —Weber, 1958 (1956). Tenth Intemat. Cong. Ent., Proc. 2:
459-473 (evolution of ants and feeding habits). — Eisner and Wilson, 1958 (1956). Tenth
Intemat. Cong. Ent., Proc. 2: 509-513 (food transmission). — KannowsM, 1959. Insectes
Sociaux 6: 115-162 (flight activities and colony founding of bog ants in Mich.). — Wilson,
1959. Anat. Rec. 134: 653 (pheromones in organization of ant societies). — Bartlett, 1961.
Ent. Soc. Amer., Ann. 54: 543-551 (influence of ants on parasites, predators, and scale
insects). — Downey, 1962. Ent. News 73: 57-66 (association with larvae of a lycaenid
butterfly). — Ayre, 1962. N. Y. Ent. Soc., Jour. 70: 159-167 (use of Lincoln Index for
estimating size of colonies). — Reid, 1962. Ga. Agr. Expt. Sta. and Univ. Ga. Handbook, 71
pp. (as intermediate hosts of chicken and turkey tapeworms). — Van Pelt, 1963. Amer.
Midland Nat. 69: 205-223 (distribution in south. Blue Ridge Mtns.). —Way, 1963. Ann. Rev.
Ent. 8: 307-344 (mutualism between ants and honeydew producing Homoptera). — Kincaid,
1963. Amer. Micros. Soc., Trans. 82: 101-105 (pollination of plants). —Wilson, 1963. Ann.
Rev. Ent. 8: 345-368 (social biology). — WUson, 1963. Evolution 17: 249-253 (social
modification). —Ayre, 1963. Canad. Ent. 95: 712-715 (feeding habits). —Wilson and
Bossert, 1963. Recent Progress 19: 673-716 (chemical communication). — Orlog, 1964 (1963).
Ent. Expt. and Appl. 6: 95-106 (plant virus transmission). — Talbot, 1965. Insectes Sociaux
12: 19-47 (populations in a low field). — Brian, 1965. Social insect populations. Academic
Press, London and N. Y., 135 pp. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326, 105 pp.
(house-infesting ants of east. U. S.). — Janzen, 1966. Evolution 20: 249-275 (coevolution of
mutualism between ants and acacias). — Beck, Allred, and Despain, 1967. Great Basin Nat.
27: 67-78 (predaceous-scavenger ants in Utah). — Sudd, 1967. An introduction to the
behavior of ants, St. Martin's Press, N. Y., 200 pp. — Markin, 1968. Jour. Econ. Ent. 61:
1744-1745 (handling techniques for large quantities of ants). — Wilson, 1971. The insect
societies, Belknap Press of Harvard Univ., 548 pp. — Lettendre, et al., 1971. Nat. Canad.
98: 591-606 (spp. from St. Hippolyte, Que.). —Payne and Mason, 1971. Ent. Soc. Wash.,
Proc. 73: 135-136, 138 (ants associated with pig carrion). — Jennings, 1971. Ent. Soc. Amer.,
Ann. 64: 384 (ants preying on jack-pine budworm larvae). — Finnegan, 1971. Canad. Ent.
103: 1489-1493 (indigenous ants as limiting agents of forest pests in Que.). — Bhatkar and
Whitcomb, 1970. Fla. Ent. 53: 229-232 (artificial diet for rearing various spp.). — Nielsson,
et al., 1971. Fla. Ent. 54: 245-248 (ants associated with aphids in Fla.). — Whitcomb, et al.,
1972. Fla. Ent. 55: 129-142 (ants of Fla. soybean fields). —Gregg, 1972. Canad. Ent. 104:
1073-1091 (northward distribution of ants in N. Amer.). — Hickman, 1974. Science 184:
1290-1292 (pollination by ants: a low-energy system). — Lettendre and Pilon, 1973. Nat.
Canad. 100: 195-235 (ant fauna of Laurentide, Que.). —Finnegan, 1974. Entomophaga 7:
53-59 (ants as predators of forest pests). — Gumey, 1975. Insect World Digest 2 (5): 19-25
(stinging ants).

Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 367^46 (thorax of wingless and
short-winged Hymenoptera). —Brown and Nutting, 1950. Amer. Ent. Soc., Trans. 75:
113-132 (wing venation). —Wilson, 1953. Quart, rev. Biol. 28: 136-156 (polymorphism).
—WUson, 1954. Insectes Sociaux 1: 75-80 (polymorphism). —Eisner, 1957. Harvard Univ.,
Mus. Comp. Zool., Bui. 116: 439-490 (studies of the proventriculus). — Glockner, 1957.
Insectes Sociaux 4: 83-90 (effect of hormones on metamorphosis). — Eisner and Brown,
1958 (1956). Tenth Intemat. Cong. Ent., Proc. 2: 503-508 (evolution and social significance
of the proventriculus). — Nachtweg, 1961. Insectes Sociaux 8: 369-381. — Tulloch, Shapiro,
and Hershenov, 1962. Brooklyn Ent. Soc., Bui. 77: 91-101 (ultrastructure of metastemal
glands). —Roth and Eisner, 1962. Ann. Rev. Ent. 7: 107-136 (chemical defenses). —Wilson,
1963. Sci. Amer. 208: 100-106 (pheromones). —Nachtweg, 1963. Insectes Sociaux 10: 43-57




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(sound organs). — Nachtweg, 1964 (1963) Insectes Sociaux 10: 359-378 (acoustics).
— Etterschank and Brown, 1964. Ent. Monthly Mag. 100: 5-7 (Malphigian tubules as
meristic characters). —Williams and Williams, 1964. Soc. Expt. Biol. Med. 116: 161-163
(toxicity studies of ant venom). —Cavil and Robertson, 1965. Science 149: 1337-1345 (ant
venoms, attractants and repellants). — Law, Wilson, and McCloskey, 1965. Science 149:
544-546 (biochemical polymorphism). —Brown, 1968. Amer. Nat. 102: 188-191 (function of
metapleural glands). — (Jotwald, 1969. Cornell Univ., Agr. Expt. Sta. Mem. 408, 150 pp.
(mouthparts). — Hermann, 1969. Ga. Ent. Soc., Jour. 4: 123-141 (poison apparatus). — Blum,
1969. Ann. Rev. Ent. 14: 57-80 (alarm pheromones).

Subfamily DORYLINAE

Members of this subfamily are known as army ants and are sometimes referred to as legiona-
ry ants in the New World and driver ants in the Old World. They are predaceous and are known
for their foraging expeditions the size of which are sometimes exaggerated. Army ants exhibit a
number of morphological and biological pectdarities not common to most ants such as wasplike
males, wingless termitelike females, blind workers, and their raiding and emigrating behavior.
Rettenmeyer (1963) outlined the following traits in which they differ from other ants: (1) They
feed almost exclusively on animal prey which is collected by large groups of raiding workers; (2)
their raiding columns usually connect to the nest by at least one continuous column; (3) the en-
tire colony periodically and frequently emigrates to new nest sites; (4) emigrations are largely
dependent on the size, caste, age, and range of ages of the brood (or broods); and (5) the colonies
are fovmded by division of an entire colony into two (or possibly several) daughter colonies.
Other ants may possess some of these traits, but not all of them.

Much of the biological work on army ants has been done in Central America on the terrestrial
species of Eciton which bivouac in large clusters above the groimd and whose colonies may
number up to a million individuals. Most of the army ants, however, are subterranean in habit,
though the raiding columns of some may appear above ground. Raiding may be in columns only
several ants wide or in swarms of a fan-shaped pattern. Most of the prey is other Arthropods,
only occasionally vertebrates. All species have nomadic and statary activity cycles where the en-
tire colony moves from one area to another, a imique behavior studied by Schneirla and Retten-
meyer in the papers listed below.

Borgmeier's revision of 1955 is the most definitive taxonomic work on this subfamily for the
New World. He recognized 137 species in 5 genera and 2 tribes. Only the tribe Ecitonini is found
in the United States; the other tribe, Cheliomyrmicini, includes a single genus of several species
foimd from Mexico to Brazil In the United States, most species are found in the Southwest,
though several reach the Atlantic coast and range north to Iowa, Ohio, and Virginia. Most of the
taxonomy is based on males and workers, and for some species only one caste is known. Further
study and association of castes may result in some synonymy, especially in Neivamyrmex.

Revision: Smith, 1942. Amer. Midland Nat. 27: 537-590 (U. S.). — Borgmeier, 1955. Studia
Ent. 3: 1-716 (New World). — Watkins, 1976. The identification and distribution of New
World army ants, Baylor Univ. Press 102 pp. (keys to genera and spp. of workers and
males; distribution maps).

Taxonomy: Wheeler, 1943. Ent. Soc. Amer., Ann. 36: 319-332 (larvae). —Borgmeier, 1953.
Studia Ent. 2: 1-51. —Borgmeier, 1958. Studia Ent. l(n. s.): 197-208 (Ecitonini;
identification of females to genus). —Wheeler and Wheeler, 1964. Ent. Soc Wash., Proc.
66: 129-137 Oarvae). —Watkins 1972. Kans. Ent. Soc., Jour. 45: 347-372 (U. S.
Neivamyrmex).

Biology: Wheeler, 1900. Amer. Nat. 34: 563-574. —Wheeler, 1910. Ants, pp. 246-266.

—Schneirla, 1933. Jour. Comp. Psychology 15: 267-299 (in Panama). —Schneirla, 1934. Nat.
Acad. ScL, Proc. 20: 316-321 (raiding and other phenomena). —Schneirla, 1938. Jour. Comp.
Psychology 25: 51-90 (theory of army ant behavior). — Schneirla, 1940. Jour. Comp.
Psychology 29: 401-460 (mass organization in the swarm-raiders). — Schneirla, 1944. N. Y.
Ent. Soc., Jour. 52: 153-192 (reproductive functions of queen as pace-makers of group
behavior). —Schneirla 1944. Amer. PhiL Soc. Proc. 87: 438-457 (nomadism in Eciton
hurchelli (Westwood)). —Schneirla, 1945. BioL BuL 88: 166-193 (nomad -statary relations in
swarmers; migration). — Schneirla, 1947. Amer. Mus. Novitates 1336: 1-20 (life and




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behavior under dry season conditions with reference to reproductive functions).
— Schneirla, 1948. Zoologica (N. Y.) 33: 89-112 (life and behavior under dry season
conditions, appearance and date of males). — Schneirla, 1948. Sci. Amer. 178: 16-23
(general). —Schneirla, 1949. Amer. Mus. Nat. Hist., Bui. 94: 1-82 (life and behavior under
dry season conditions, course of reproduction and colony behavior). — Schneirla, 1950.
(Mex.) Inst, de Biol., An. 20: 371-384 (environmental adaptations). — Schneirla and Brown,
1950. Amer. Mus. Nat. Hist., Bui. 95: 269-233 (life and behavior under dry season
conditions, cyclic processes in behavioral and reproductive functions). — Schneirla and
Brown, 1952. Zoologica (N. Y. ) 37: 5-32 (sexual broods and production of young queens).
— Schneirla, 1953. Insectes Sociaux 1: 29-41 (army ant queen). — Schneirla, Brown and
Brown, 1954. Ecol. Monog. 24: 269-296. (bivouac or temporary nest as adaptive factor in
terrestrial species). — Tafuri, 1955. N. Y. Ent. Soc, Jour. 63: 21-41 (growth and
polymorphism of larva of Eciton hamatum (F.)). — Schneirla, 1956. Insectes Sociaux 3:
49-69 (colony division and related processes). — Schneirla, 1956. Smithsn. Inst. Ann. Rpt.,
1955 pp. 379-406 (army ants, general). —Schneirla, 1957. Insectes Sociaux 4: 259-298
(comparison of functional patterns in army ants). — Schneirla, 1957. Amer. Phil. Soc, Proc.
101: 106-133 (theoretical consideration of cyclic processes). — Schneirla, 1958. Insectes
Sociaux 5: 215-255 (behavior and biology of Neivamyrmex nigrescens (Cresson) and N.
opacithorax (Emery)). — Brown, 1960. Psyche (56: 25-27 (alarm and attack behavior).
— Schneirla, 1960. Smithsn. Treasury of Science. Simon and Schuster, N. Y., pp. (564-696
(army ants, general). —Schneirla, 1961. Ztschr. f. Tierpsychol. 18: 1-32 (sexual broods and
colony division of Neivamyrmex nigrescens (Cresson)). — Rettenmeyer, 1961. Kans. Univ.,
Sci. Bui. 42: 993-1066 (biology and taxonomy of flies over swarms of army ants).
—Rettenmeyer, 1963. Univ. Kans., Sci. Bui. 44: 287-465 (behavioral studies of army ants in
Canal Zone and Kansas). —Schneirla, 1963. Animal Behavior 11: 583-595 (springtime
resurgence of cyclic function, Arizona species). — Rettenmeyer, 1960. XI. Intemat. Kong,
fur Ent., Wien. 1960, 1: 610-612 (mites associated with army ants). —Rettenmeyer, 1962.
Kans. Ent. Soc., Jour. 35: 377-384 (millipedes associated with army ants). —Rettenmeyer,

1963. Ent. Soc. Amer., Ann. 56: 170-174 (Thysanura associated vdth army ants). — Watkins,

1964. Kans. Ent. Soc, Jour. 37: 22-28 (trail following). —Watkins and Cole, 1966. Tex. Jour.
Sci. 18: 254-265 (attraction of workers to secretion of queens). — Akre and Rettenmeyer,
1966. Kans. Ent. Soc, Jour. 39: 745-782 (Staphylinidae associated vdth army ants).
—Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc, Jour. 40: 146-151 (trail follov^dng and
trail preference). —Akre and Rettenmeyer, 1968. Kans. Ent. Soc, Jour. 41: 165-174 (trail
following by guests of army ants). —Akre, 1968. Pan-Pacific Ent. 44: 87-101 (Histeridae
associated with army ants). — Rettenmeyer and Akre, 1968. Ent. Soc Amer., Ann. 61:
1317-1326 (ectosymbiosis between phorid flies and army ants.). — Kannowsld, 1969.
Intemat. Union for study of social insects, VI Cong., Proc 6: 77-83 (daUy and seasonal
periodicities in nuptial flights). — Torgerson and Akre, 1970. Melanderia 5: 1-28
(persistence of army ant chemical trails and their significance in Ecitonine-Ecitophile
association). —Torgerson and Akre, 1970. Kans. Ent. Soc, Jour. 43: 395-404 (interspecific
responses to trail and alarm pheromones). —Schneirla, 1971. Army Ants: A Study in Social
Organization. W. H. Freeman, San Francisco. 349 pp. — Topoff, 1971. Amer. Nat. 105:
529-548 (polymorphism related to division of labor and colony cyclic behavior).

Morphology: Hagen, 1954. Amer. Mus. Novitates 1663: 1-12 (anatomy of queen of Eciton).
— Hagen, 1954. Amer. Mus. Novitates 1664: 1-17 (Reproductive system of queen of
Eciton). —Hagen, 1954. Amer. Mus. Novitates 1665: 1-20 (oocyte cycle). — Lappano, 1958.
Insectes Sociaux 5: 31-66 (morphological study of larval development in all worker broods
of Eciton burchelli (Westwood). — Whelden, 1963. N. Y. Ent. Soc, Jour. 71: 158-178
(reproductive system of worker and female of Eciton hxiniatum{F.) and Eciton burchelli
(Westwood)). —Whelden, 1963. N. Y. Ent. Soc, Jour. 71: 246-261 (antennae and legs of £.
hamatum and E. burchelli).

Tribe ECITONINI

Genus LABIDUS Jurine

Labidus Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 282.
Type-species: Labidus latreillii Jurine. Monotypic




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Nycteresia Roger, 1861. Berlin. Ent. Ztschr. 5: 21.

Type-species: Formica coeca Latreille. Monotypic.
Pseudodicthadia Andre, 1885. Spec. Hym. Eur. Alg. 2: 838.

Type-species: Pseudodicthadia iyicerta Andre. Monotypic.

A neotropical genus of 8 species, one of which reaches the United States.

Revision: Borgmeier, 1955. Studia Ent. 3: 80-134.

Taxonomy: Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc. 66: 134 (larvae).

Biology: Rettenmeyer, 1963. Kans. Univ., Sci. Bui. 44: 403-424 (behavioral studies in Canal

Zone).

Labidus coecus (Latreille)
S. Ark., La., Okla., Tex. s. to Argentina. Ecology: The large colonies, usually
with many thousands of individuals, are found in more or less temporary nests in
decayed logs and stumps or in ground beneath objects. They are subterranean and
nocturnal. Food consists of other arthropods, small mammals, birds, and nuts. Workers
are highly predaceous and are known to feed on injurious insects such as the immature
stages of the screwworm (Cochliomyia hominivorax (Coquerel)) and the secondary
screwworm (C. macellaria (F.)).

Formica omnivora Olivier, 1791. Encycl. Meth. Hist. Nat. 6: 496. ♀. Preocc. by Linnaeus,
1758.

Formica coeca Latreille, 1802. Hist. Nat. Fourmis, v. 9, p. 270. ♀.

Labidus latreillii J urine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 283. ♂.

Labidus jurini Shuckard, 1840. Ann. Nat. Hist. 5: 198. ♂.

Labidus servillei Westwood, 1842. Arcana Ent., v. 1, p. 75. ♂.

Mutilla (Labidus) fulvescens Blanchard, 1849. In Cuvier, Regne Animale, ed. 3, v. 2, pi.
118, fig. 2.

Labidus saji (!) Haldeman, 1852. In Stanbury, Exped. Great Salt Lake, p. 367. ♂.

Labidus atriceps Smith, 1859. Cat. Hym. Brit. Mus., v. 7, p. 5. ♂.

Labidus pilosus Smith, 1859. Cat. Hym. Brit. Mus., v. 7, p. 7. ♂.

Labidus panzeri Smith, 1859. Cat. Hym. Brit. Mus., v. 7, p. 72. ♂.

Eciton vastator Smith, 1860. Jour. Ent., London 1: 71. ♂.

Eciton erratica Smith, 1860. Jour. Ent., London 1: 71. ♂.

Myrmica rubra Buckley, 1867. Ent. Soc. Phila., Proc. 6: 335. ♂. Preocc. in Myrmica by
Linnaeus, 1758.

Pseudodicthadia incerta Andre, 1885. Spec. Hym. Eur. Alg. 2: 838. ♀.

Eciton smithii Dalla Torre, 1892. Cat. Hym., v. 7, p. 6. N. name for L. pilosus Smith.

Eciton coecum var. biloba Emery, 1901. Soc. Ent. Belg., Ann. 45: 51. ♂.

Eciton nigrita Emery, 1901. Soc. Ent. Belg., Ann. 45: 52. ♂.

Eciton coecum var. kulowi Forel, 1901. Mitt. Nat. Mus. Hamburg 18: 47. ♂.

Eciton selysi Forel, 1904. Soc. Ent. Belg., Ann. 48: 169. ♂.

Eciton grassator Forel, 1911. Deut. Ent. Ztschr., p. 288. ♀.

Eciton (Labidus) coecum servillei var. hostilis Santschi, 1920. Soc. Ent. France, Ann. 88:
368. ♂.

Eciton (Labidus) coecum var. opcunfrons Wheeler, 1921. Amer. Acad. Arts and Sci., Proc.
56:310. ♂.

Eciton (Labidus) coecum var. elsbethae Forel, 1922. Rev. Suisse Zool. 30: 91. ♂.

Eciton (Labidus) serpentis Weber, 1938. Ent. Soc. Amer., Ann. 31: 209. ♂.

Taxonomy: Weber, 1941. Amer. Midland Nat. 26: 238 (queen). —Wheeler, 1943. Ent. Soc.
Amer., Ann. 36: 332 Oarva). —Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc. 66: 135
(larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., BuL 24: 408-409. — Lindquist, 1942. Jour.
Econ. Ent. 35: 850 (as predators of screwworms). —Hess, 1958. Field and Lab. 26: 35-37.
— Kempf, 1961. Studia Ent. 4(n.s.): 551-552 (as a cave ant). — Rettenmeyer, 1963. Kans.
Univ., Sci. Bui. 44: 41&424. —Smith, 1965. U. S. Dept. Agr. Tech. Bui. 1326, pp. 17-18.
— Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-26i5 (attraction of workers to secretion of
queens). —Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail
following and trail preference).







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Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 390 (maxillary and labial palps).
— Gotwald, 1969. N. Y. Agr. Expt. Sta. (Cornell Univ.) Mem. 408, p. 127 (mouthparts).

Genus NOMAMYRMEX Borgmeier

Eciton subg. Nomamyrmex Borgmeier, 1936. Inst. Biol. Veg. Arq. 3: 55.
Type-species: Eciton crassicome Smith. Orig. desig.

A genus of two species, a subspecies of only one reaching the United States. They are subter-
ranean though carrying on terrestrial raids. Raiding is in columns of several ants wade and a few
meters long and is sometimes done during the daylight hours.

Revision: Borgmeier, 1955. Studia Ent. 3: 135-161.

Taxonomy: Borgmeier, 1953. Studia Ent. 2: 4. —Borgmeier, 1958. Studia Ent. 1 (n. s.):
201-203.

Biology: Rettenmeyer, 1963. Kans. Univ. Sci. Bui. 44: 424-432 (behavioral studies in Canal
Zone; esenbeckii crassicomis (Smith)).

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 390 (maxillary and labial palps).
esenbeckii wilsoni (Santschi). Extreme s. Tex. to Costa Rica. N. esenbeckii esenbeckii

(Westwood) is South American. Literature references for esenbeckii (Westwood) and
crassicomis (Smith) from Texas pertain to wilsoni.
Eciton {Labidits) Esenbecki Wilsoni Santschi, 1919. Soc. Ent. France, Ann. 88: 366. ♂.
Eciton (Holopone) crassicomis mordax Santschi, 1928. Deut. Ent. Ztschr., p. 415. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 409 (male).

Genus NEIVAMYRMEX Borgmeier ^ *

Eciton subg. Acamatus Emery, 1894. Soc. Ent. Ital., Bol. 26: 181. Preocc. by Schoenherr,
1833.

Type-species: Eciton {Acamatus) schmitti Emery. Desig. by Wheeler, 1911.
Eciton subg. Neivamyrmex Borgmeier, 1940. Rev. de Ent. 11: 606. N. name for Acamatiis

Emery.
Woitkowskia Enzmann, 1952. Iowa Acad. Sci., Proc. 59: 443.

Type-species: Woitkowskia connectens Enzmann. Orig. desig.

About 115 species are known for this New World genus, but only 23 are found north of Mex-
ico. Most species are found in the southwestern states with a few ranging east to the Atlantic
coast and as far north as Virginia, Ohio, Illinois, and Iowa. The species are hypogaeic and are
found in the soil under objects, though some have been reported from rotten logs and stumps.
Some may carry on their foraging and emigrating activities during daylight, but most are ap-
parently nocturnal in their activities. Most colonies have one functional queen, and new colonies
are produced by fission. The cyclic pattern of nomadic and statary phases is similar to the tropi-
cal species, but in most Nearctic species the activity stops in the autumn and resumes again in
the spring. Because many of the species listed below were described from a single caste, future
study and caste association may reveal that fewer species actuaUy exist.

Revision: Smith, 1942. Amer. Midland Nat. 27: 537-590 (U. S. species). —Borgmeier, 1955.
Studia Ent. 3: 277-651 (New World species).

Taxonomy: Borgmeier, 1950. Rev. de Ent. 21: 624. — Watkins, 1971. Kans. Ent. Soc., Jour. 44:
93-103 (key to major workers and queens of U. S.). —Watkins, 1972. Kans. Ent. Soc., Jour.
45: 347-372 (keys for each caste and distribution of U. S. species).

Biology: Schneirla, 1958. Insectes Sociaux 5: 215-255 Oast part of fimctional season, s.e.
Ariz.). —Schneirla, 1961. Ztschr. f. Tierpsychologie 18: 1-32 (sexual broods and colony
division). —Schneirla, 1963. Animal Behaviour 11: 583-595 (spring resurgence of cyclic
function, s.e. Ariz.). —Rettenmeyer, 1963. Kans. Univ., Sci. Bui. 44: 433-452 (behavioral
studies). —Watkins, 1964. Kans. Ent. Soc., Jour. 37: 22-28 (trail foUowing). —Watkins and
Cole, 1966. Tex. Jour. Sci. 18: 254-265 (attraction of workers to secretions of queens).
—Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail following and




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trail preference). — Plsek, Kroll, and Watkins, 1969. Kans. Ent. See., Jour. 42: 452-456
(association with carabids).

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 392-393 (maxillary and labial palps).
ag^lis Borgmeier. S. Ariz.; Mexico. Only the worker is known.

Neivamyrmex agilis Borgmeier, 1953. Studia Ent. 2: 45. ♂.

Neivamyrmex andrei (Emery)
N. Mex., Ariz.; Mexico. Only the male is known.

Eciton andrei Emery, 1901. Soc. Ent. Ital., Bol. 33: 53. ♂.

Eciton (Acamatus) oslari Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 415. ♂.

Neivamyrmex baylori Watkins
Tex. (Waco).

Neivamyrmex baylori Watkins, 1973. Kans. Ent. Soc., Jour. 46: 430-433. ♂.
califomicus (Mayr). Utah, Nev., Calif. Only the worker and queen are known.

Eciton califamicum Mayr, 1870. Zool.-Bot. Gesell. Wien., Verb. 20: 969. ♂.

Eciton (Acamatus) califomicum. var. obscura Forel, 1914. Soc. Vaud. Sci. Nat., Bui. 50:
265. ♂.

Taxonomy: Watkins, 1972. Kans. Ent. Soc., Jour. 45: 363-366 (queen, distribution).

Biology: Mallis, 1938. Sci. Monthly 47: 220-226. — Mallis, 1941. South. Calif. Acad. Sci., Bui.
40: 62.

Neivamyrmex carolinensis (Emery)
Va., N. C, S. C, Ga., Fla., Ohio, Tenn., Ala., Miss., La., Nebr., Kans., N.
Mex., Ariz. All castes are known.
Eciton (Acamatus) carolinense Emery, 1894. Soc. Ent. Ital., Bol. 26: 184. ♂.

Biology: Smith, 1928. Ent. News 39: 245. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 278, 304.
—Watkins, 1964. Kans. Ent. Soc., Jour. 37: 22-28 (trail following). —Watkins and
Rettenmeyer, 1967. Psyche 74: 228-233 (effect of queen on longevity of workers).

Neivamyrmex fallax Borgmeier
La., Kans., Tex., N. Mex., Ariz.; Mexico; Guatemala. Only the worker is

known. Literature references for commutatum (Emery) for N. Amer. north of Mexico
should be referred to fallcuc.
Neivamyrmex fallax Borgmeier, 1953. Studia Ent. 2: 48. ♀.

Neivamyrmex fuscipennis (Wheeler)
Kans., e. Tex. Only the male is known.

Acamatus fiiscipennis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 417. 6. Misdet. as
spoliator Forel, hut fiiscipennis validated by pi. 26, fig. 12.

Taxonomy: Watkins, 1975. Southwest. Nat. 20: 85-90 (relationship to N. macropterus
Borgmeier; lectotype).

Neivamyrmex harrisii (Haldeman)
Okla., Tex., N. Mex., Ariz.; Mexico. AH castes are known. Records citing
this species from Utah are probably incorrect.
Labidus harrisii Haldeman, 1852. In Stanbury, Exped. Great Salt Lake, p. 367. ♂.
Eciton wheeleri Emery, 1901. Soc. Ent. Ital., Bol. 33: 65. ♂.
Eciton (Acamutu^) wheeleri dubia Creighton, 1932. Psyche 39: 75. 9,9.

Taxonomy: Watkins, 1968. Amer. Midland Nat. 80: 273-275 (association of castes).
—Borgmeier, 1958. Studia Ent. 1: 206-207.

Biology: Watkins and Cole, 1966. Tex. Jour. Sci 12: 254-265 (attraction of workers to
secretion of queens). — Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40:
146-151 (trail following and trail preference). —Plsek, Kroll, and Watkins, 1969. Kans. Ent.
Soc., Jour. 42: 452-456 (carabids in raiding columns).

Morphology: Forbes and Do-Van-Quy, 1965. N. Y. Ent. Soc., Jour. 73: 95-111 (nude
reproductive system).

Neivamyrmex leonardi (Wheeler)
Okla., Tex., Calif.; Mexico. Only the worker is known.

Eciton (Acamatus) leonardi Wheeler, 1915. Amer. Mus. Nat. Hist., BuL 34: 392. ♀.
Eciton (Acamatus) peninsulare Mann, 1926. Psyche 33: 98. ♂.

Taxonomy: Borgmeier, 1953. Studia Ent. 2: 10. —Watkins, 1971. Kans. Ent. Soc., Jour. 44:
101-103 (worker; biological notes).




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Biology: Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-265 (=pauxillus of authors, not *
Wheeler; attraction of workers to secretion of queens). —Watkins, Cole, and Baldridge,
1967. Kans. Ent. Soc, Jour. 40: 146-151 (=pauxillus of authors, not Wheeler; trail
following and trail preference).

Neivamyrmex macropterus Borgmeier
W. Tex., N. Mex., Ariz.; Mexico, only the male is known.
Neivamyrmex macropterus Borgmeier, 1953. Studia Ent. 2: 40. ♂.

Taxonomy: Watkins, 1975. Southwest. Nat. 20: 85-89 (relationship to fuscipennis (Wheeler)).

Neivamyrmex melanocephalus (Emery)
S. Ariz, to Honduras. Only the worker is known.

Eciton (AcamatiLs) melanocephalum Emery, 1895. Zool. Jahrb. Abt. f. System. 8: 260. ♂.
Eciton {Acamatv^) melanocephalum xipe Wheeler, 1914. N. Y. Ent. Soc., Jour. 22: 41. ♀.

Neivamyrmex melsheimeri (Haldeman)
La., Okla., Tex.; s. to Costa Rica. Only the male is known. Records
citing this species from Utah are probably incorrect.
Labidus melshaemeriH) Haldeman, 1852. In Stanbury, Exped. Great Salt Lake, p. 368. ♂.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 418.

Neivamyrmex microps Borgmeier
Ariz. (Phoenix). Only the male is known.

Neivamyrmex microps Borgmeier, 1955. Studia Ent. 3: 635. ♂.

Neivamyrmex minor (Cresson)
Kans., Okla., Tex., N. Mex., Ariz., Nev., Calif.; Mexico. Only the male is
known. One of the smallest of our legionary ants.
Labidus m,inor Cresson, 1872. Amer. Ent. Soc., Trans. 4: 195. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 418 (male).

Biology: Wheeler and Long, 1901. Amer. Nat. 35: 165. —Cole, 1966. Brigham Young Univ.,
Sci. Bui. 7: 26.

Neivamyrmex mojave (Smith)
Calif. (Mojave Desert). Only the male is known.

Eciton (.Neivamyrmex) mojave Smith, 1943. Lloydia 6: 196. ♂.

Neivamyrmex moseri Watkins
La., Tex. Ecology: Collected from a nest of Atta texana (Buckley) in La. Only
the worker and queen are known.
Neivamyrmex moseri Watkins, 1968. Kans. Ent. Soc., Jour. 41: 528-531. ♂.

Taxonomy: Watkins, 1971. Kans. Ent. Soc., Jour. 44: 95-99 (worker, queen; also biological

notes).

Neivamyrmex nigrescens (Cresson)
W. Va., Ky., Tenn., Ga., Ala., Miss., 111., Iowa, Mo., Ark., La., Nebr., Kans.,
Okla., Tex., Colo., N. Mex., Ariz., Calif.; Mexico. Ecology: Temporary nesting sites are in
decayed logs or stimips or in the ground beneath stones and other objects. Many
foraging activities take place in daylight, and they are highly predaceous on other
insects. Large colonies contain 150,000 to 250,000 workers. Each colony has one
functional queen, and new colonies are formed by splitting. All castes are known.

Labidus nigrescens Cresson, 1872. Amer. Ent. Soc., Trans. 4: 194. ♂.

Eciton (Acamatus) Schmitti Emery, 1894. Soc. Ent. Ital, Bol. 26: 183. ♂.

Taxonomy: Wheeler, 1943. Ent. Soc. Amer., Ann. 36: 331 (larva). —Watkins, 1972. Kans. Ent.
Soc., Jour. 45: 358-363 (worker, distribution).

Biology: Wheeler, 1900. Amer. Nat. 34: 565-574 (sumichrasti, not Norton). —Smith, 1927.
Ent. Soc. Amer., Ann. 20: 401-404. —Cole, 1953. Tenn. Acad. Sci., Jour. 28: 84. — Schneirla,
1958. Insectes Sociaux 5: 215-255. — Rettenmeyer, 1963. Univ. Kans., Sci. Bui. 44: 44fr452.
—Gregg, 1963. Ants of Colo., pp. 286-288. —Watkins, 1964. Kans. Ent. Soc., Jour. 37: 22-28
(trail following). —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 18-19. —Watkins and
Cole, 1966. Tex. Jour. Sci. 18: 254-265 (attraction of workers to secretion of queens).
—Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail following and
trail preference). —Watkins, (Jehlbach, and Baldridge, 1967. Southwest. Nat. 12: 455-462
(blind snake follows pheromone trails of nigrescens). —Watkins, Gehlbach, and KroU,
1969. Ecology 50: 1098-1102 (attractant, repellant secretions). — Plsek, Kroll, and Watkins,
1969. Kans. Ent. Soc., Jour. 42: 452-456 (carabids in raiding columns). — Topoff, 1969. N. Y.
Ent. Soc., Jour. 77: 273-274 (communication). —Topoff, 1969. Psyche 76: 375-381 (predatory
association between carabids and nigrescens). —Topoff, 1970. N. Y. Ent. Soc., Jour. 78:
239-240 (cyclic behavior). —Watkins, Gehlbach, and Plsek, 1972. Tex. Jour. Sci. 23: 34.
(behavior of blind snakes in response to raiding columns).




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Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 393 (maxillary and labial palpi).

Neivamyrmex opacithorax (Emery)
Va., N. C, S. C, Ga., Fla., Tenn., Ala., Mo., Ark., Kans., Okla., Tex., N.
Mex., Ariz., Calif.; Mexico to Costa Rica. Ecology: Habits are similar to those of

Neivamyrmex nigrescens (Cresson)
All castes are known.
Eciton (Acamatus) califomicum opacithorax Emery, 1894. Soc. Ent. Ital., Bol. 26: 184. ♂.
Eciton {Acamatus) opacithorax castaneum Borgmeier, 1939. Rev. de Ent. 10: 416. ♀.

Biology: Wheeler and Long, 1901. Amer. Nat. 35: 163, 172. —Smith, 1924. Ent. News 35: 84.
—Cole, 1953. Tenn. Acad. Sci., Jour. 28: 84. — Schneirla, 1958. Insectes Sociaux 5: 215-255.
— Rettenmeyer, 1963. Univ. Kans., Sci. Bui. 44: 437-446. — Watkins, 1964. Kans. Ent. Soc.,
Jour. 37: 22-28 (trail foUowing). —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 19-20.
^Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-2(55 (attraction of workers to secretion of
queens). —Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc, Jour. 40: 146-151 (trail
following and trail preference). — Watkins, (jehlbach, and Baldridge, 1967. Southwest. Nat.
12: 455-462 (a blind snake follows pheromone trails of opicithorax).

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 393 (maxillary and labial palpi).

Neivamyrmex pauxillus (Wheeler)
Tex.; Mexico. Only the worker and queen are known.
Eciton (Acamatus) pauxillum, Wheeler, 1903. Psyche 10: 93. ♂.

Taxonomy: Watkins, 1971. Kans. Ent. Soc., Jour. 44: 99-101 (worker, queen; biological notes).

Biology: Smith, 1938. Ent. Soc. Wash., Proc. 40: 158.
pilosus mandibularis (Smith). N. Mex., Ariz. Only the male is known. N. pilosus pilosus
(Smith) is found in S. Amer.
Eciton (Neivamyrmex) pilosum mandihulare Smith, 1942. Amer. Midland Nat. 27: 543,
548. ♂.
pilosus mexicanus (Smith). Miss., Ark., La., Okla., Tex., Calif., s. to Colombia. AU castes are
known.
Labidus Mexicanus Smith, 1859. Cat. Hym. Brit. Mus., v.7, p. 7. ♂.
Eciton clavicomis Norton, 1868. Amer. Ent. Soc., Trans. 2: 46. ♀.
Eciton (Labidus) subsulcatum Mayr, 1886. Zool.-Bot. (Jesell. Wien, Verh. 36: 440. ♂.
Eciton (Acamatiis) aztecum Forel, 1901. Mitt. Nat. Mus. Hamburg 18: 49. ♂.
Eciton (Labidus) spininode militarium Santschi, 1929. Wien Ent. Ztg. 46: 85. ♂.

Taxonomy: Borgmeier, 1936. Inst. Biol. Veg. Arq. 3: 60. — Reichensperger, 1939. Zool. Jahrb.
Abt. f. System. 73: 297-300.

Biology: Smith, 1924. Ent. News 35: 85. —Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-265
(attraction of workers to secretion of queens). — Watkins, Cole, and Baldridge, 1967. Kans.
Ent. Soc, Jour. 40: 146-151 (trail foUowing and trail preference).

Neivamyrmex rugulosus Borgmeier
S. Ariz.; Mexico. Only the worker is known.

Neivamyrmex rugulosus Borgmeier, 1953. Studia Ent. 2: 49. § .

Neivamyrmex swainsonii (Shuckard)
La., Tex., N. Mex., Ariz., Calif.; s. to Argentina. Only the male is
known.
Labidus swainsonii Shuckard, 1840. Ann. Nat. Hist. 5: 201. ♂.

Eciton (Acamatus) arizonense Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 414. ♂.

Neivamyrmex texanus Watkins
Va., N. C, S. C, Ga., Fla., Tex., Colo., N. Mex., Ariz.; Mexico. All castes are
known.
Neivamyrmex texanus Watkins, 1972. Kans. Ent. Soc, Jour. 45: 353-358. 5 , 9 , 6 .

Taxonomy: Wheeler and Long, 1901. Amer. Nat. 35: 161 (male described as schmitti).
— Borgmeier, 1955. Studia Ent. 3: 496 (male described as nigrescens).

Unplaced Taxon of Doryunae

Myrmica (Mofrwmarium(!)) coeca Buckley, 1867. Ent. Soc. Phila., Proc. 6: 339. 5. Tex. (San
Saba Co.).




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Subfamily CERAPACHYINAE

This small subfamily, found in the tropical regions of the world, exhibits a blending of doryline
and ponerine traits both morphologically and biologically. Only three species are known from
north of Mexico. Little is known concerning the behavior of the New World forms, but they are
predaceous and carnivorous and the colonies are small. Wilson (1958) studied the behavior of
several species from Melanesia and Australia and found them all to be myrmecophagous, feeding
on the broods and sometimes adults of other species of ants. He suggested that these ants carry
on an alternating, group foraging and raiding behavior pattern by which the colony efficiently
exploits the surrounding territory.

After this section was completed, Brown (1975) published on the Cerapachyinae and regarded
the Cerapachyini and Acanthostichini as tribes of the Ponerinae, consequently not recognizing
the Cerapachyinae as a subfamily.

Revision: Brown, 1975. Search, Agr., Ent. (Ithaca) 15, 5 (1): 14-36 (Cerapachyini and
Acanthostichini as tribes of Ponerinae; biol. notes; keys to world genera and species).

Taxonomy: Wheeler, 1920. Psyche 27: 50-51. — Donisthorpe, 1921. London Ent. Soc., Proc.,
pp. xlv-xlvii. —Wheeler, 1922. Amer. Mus. Nat. Hist., Bui. 45: 51-52. — Morley, 1939. Soc.
Ent. France, Bui. 44: 114-118. — Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 421-422.
— Creighton, 1950. Harvard Univ., Mus. Compar. Zool., Bui. 104: 56-59. —Wheeler, 1950.
Psyche 57: 102-113 (larvae). — Kuznezov, 1952. Dusenia 3: 115. —Brown, 1954. Insectes
Sociaux 1: 26-27. —Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc. 66: 65-71 (larvae).
—Wheeler and Wheeler, 1973. Psyche 80: 204-211 (larvae).

Biology: Wilson, 1958. Insectes Sociaux 5: 129-140 (behavior of certain Melanesian and
Australian species).

Tribe CERAPACHYINI

Genus CERAPACHYS Smith

Cerapachys Smith, 1858. Linn. Soc. London, Jour. 2: 74.

Type-species: Cerapachys antennatus Smith. Monotypic.
Syscia Roger, 1861. Berlin. Ent. Ztschr. 5: 19.

Type-species: Syscia typhla Roger. Monotypic.
Parasyscia Emery, 1882. In Andre, Spec. Hym. Eur. Alg. 2: 235.

Type-species: Parasyscia piochardi Emery. Monotypic.

A pantropical genus with about 8 New World species, two of which are found in southwestern
United States. For additional generic synonymy, see Brown, 1975 Search, Agr., Ent. (Ithaca) 15,
5 (1): 18-19.

Taxonomy: Kempf, 1972. Studia Ent. 15: 76 (generic synonymy).

Cerapachys augustae Wheeler
Tex., Ariz.; Mexico. The type colony from Austin, Tex., was found 6 inches
below the surface of soil containing limestone chips; 10 workers and a female were
found. Other specimens were foimd in the stomach of an armadillo.
Cerapachys (Parasyscia) augustae Wheeler, 1902. Biol. Bui. 3: 182. ♂, ♀.

Taxonomy: Smith, 1942. Ent. Soc. Wash., Proc. 44: 63 (male). —Wheeler, 1950. Psyche 57:
106-107 (larva).

Biology: Wheeler, 1903. Psyche 10: 205-209.

Cerapachys davisi Smith
Tex. (Ft. Davis, 5,000 ft.).

Cerapachys (Parasyscia) davisi Smith, 1942. Ent. Soc. Wash., Proc. 44: 64. ♂.

Tribe ACANTHOSTICHINI

Genus CTENOPYGA Ashmead

Ctenopyga Ashmead, 1905. Canad. Ent. 37: 382. Nom. nud.
Ctencypyga Ashmead, 1906. Ent. Soc. Wash., Proc. 8: 29.

Type-species: Ctenopyga tottmsendi Ashmead. Monotypic.




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Ctenopyga texanus (Forel)
Tex.; Mexico.

Acanthostichiis texanus Forel, 1904. Soc. Ent. Belg., Ann. 48: 168. ♀.
Ctenopyga tawnsendi Ashmead, 1906. Ent. Soc. Wash., Proc. 8: 29-30. ♀, ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 400 (female). —Smith, 1955.
Brooklyn Ent. Soc, Bui. 50: 48-50 (synonymy).

Subfamily PONERINAE

One of the smaller subfamilies of ants, with most species found in the tropical regions of the
world. Most of the North American forms are either northern extensions of Neotropical taxa or
have been introduced by commerce from other parts of the world. These are primitive ants
which nest in small colonies of a few hundred individuals or less, mostly in soil or rotting wood.
They are predaceous and carnivorous.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 111-144 (larvae). —Wheeler
and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 443-462 (larvae). —Wheeler and Wheeler,
1971. Ent. Soc. Amer., Ann. 64: 1197-1217 (larvae). —Wheeler and Wheeler, 1974. Ent. Soc.
Wash., Proc. 76: 278-281 (larvae). —Wheeler and Wheeler, 1976. Amer. Ent. Soc, Trans.
102: 41-64 (supplementary studies on larvae).

Biology: Wheeler, 1910. Ants, pp. 225-245. — WUson, 1958. Evolution 12: 24-31 (beginnings of
nomadic and group-predatory behavior).

Tribe AMBLYOPONINI

Revision: Brown, 1960. Harvard Univ., Mus. Comp. ZooL, Bui. 122: 145-230.

Genus AMBLYOPONE Erichson

Amblyopone Erichson, 1842. Arch. f. Naturgesch. 8: 260.

Type-Sf>ecies: Amblyopone australis Erichson. Monotypic.
Stigmatornma Roger, 1859. Berlin Ent. Ztschr. 3: 250.

Type-species: Stigmatornma denticulatum Roger. Desig. by Bingham, 1903.
Arotropus Provancher, 1881. Nat. Canad. 12: 205.

Type-species: Arotropus binodostis Provancher. Monotypic.
Amblyopopana Schulz, 1906. Spolia Hym., p. 154. Emend.
Stigmatmnma subg. Xymmer Santschi, 1914. Bol. Lab. Zool. Gen. e Agr. Portici 8: 311.

Type-sf)ecies: Stigmatornma {Xymmer) muticum Santschi. Monotypic.
Stigmatornma subg. Fulakora Mann, 1919. Harvard Univ., Mus. Comp. Zool., Bui. 63: 279.

Type-sp»ecies: Stigmatornma {Fulakora) celata Mann. Orig. desig.
Amblyopone subg. Neoamblyopone Clark, 1927. In Wheeler, Amer. Acad. Arts and Sci.,
Proc 62: 1.

Type-species: Amlyopone {Neoamblyopone) clarki Wheeler. Monotypic.
Amblyopone subg. Protamblyopone Clark, 1927. In Wheeler, Amer. Acad. Arts and Sci.,
Proc. 62: 1.

Type-species: Amblyopone {Protamblyopone) aberrans Wheeler. Monotypic.
Lithomyrmex Clark, 1928. Roy. Soc. W. Austrjilia, Jour. 14: 30.

Type-si)ecies: Lithomyrmex gUvuerti Clark. Orig. desig.
Ericapelta Kusnezov, 1955. Zool. Anz. 154: 273.

Type-species: Ericapelta egregia Kusnezov, Monotypic.

This genus is represented in the tropical and temperate regions of the world though it is more
highly developed in the Australian Region than elsewhere. The Nearctic forms commonly occur
in wooded areas, especially those that are well shaded. They are subterranean, and the workers
are timid and slow of movement. The female of pallipes forages for food during the period of
nest founding, an archaic habit in ants.

Revision: Creighton, 1940. Amer. Mus. Novitates 1079: 1-8. —Brown, 1960. Harvard Univ.,
Mus. Comp. Zool., BuL 122: 155-169.

Taxonomy: Brown, 1949. Psyche 56: 81-88. —Wheeler and Wheeler, 1964. Ent. Soc. Amer.,
Ann. 57: 444-446 (larvae).




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Biology: Haskins and Haskins, 1951. Amer. Midland Nat. 45: 432-445 (colony founding of A.
aiistralis Erichson).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. ZooL, Bui. 116: 476 (proventriculus).

Amblyopone oregonense
***authority mismatch
CWheeler). B. C, Wash., Oreg., Calif. Ecology: Usually at low elevations in the
coastal mountains.
Stigmatomma pallipes oregonense Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 389. ♂.
9.
Taxonomy: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bui. 122: 169, 183.

Amblyopone pallipes (Haldeman)
Ont., Que. s. to Ga. w. to Wis., Iowa, Okla., Colo., Tex., Ariz. Ecology: The
small colonies are most common in areas of heavy cover and considerable precipitation.
Chilopods appear to be the main food source.
Typhlopone pallipes Haldeman, 1844. Acad. Nat. Sci. Phila., Proc. 2: 54. ♂.
Stigmatomma serratum Roger, 1859. Berlin. Ent. Ztschr. 3: 251. ♂.
Arotrojms binodosus Provancher, 1881. Nat. Canad. 12: 206. ♀.
Stigmatomma pallidipesC.) var. Wheelen Santschi, 1913. Soc. Ent. Belg., Ann. 57: 429. ♂.

9,3.
Stigmntomma pallipes arizonense Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 389. ♂.
Stigmatomma pallipes m,ontigena Creighton, 1940. Amer. Mus. Novitates 1079: 2, 7. ♂, ♀.
Stigmatomma pallipes subterranea Creighton, 1940. Amer. Mus. Novitates 1079: 3, 8. ♂.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 261-262 (each caste). —Wheeler and
Wheeler, 1952. Amer. Midland Nat. 48: 114 (larva). — Brovm, 1960. Harvard Univ., Mus.
Comp. Zool, Bui. 122: 169, 183-185. — Francoeur and Beique, 1966. Canad. Ent. 98: 141
(Provancher types).

Biology: Wheeler, 1900. Biol. Bui. 2: 56-64. —Haskins, 1928. N. Y. Ent. Soc., Jour. 36: 179-184.
— Buren, 1944. Iowa State Col., Jour. Sci. 18: 279. —Cole, 1953. Tenn. Acad. Sci., Jour. 28:
84. —Gregg, 1963. Ants of Colo., pp. 279-282.

Morphology: Whelden, 1957. N. Y. Ent. Soc., Jour. 65: 1-21 (anatomy). — Gotwald, 1969. N. Y.
Agr. Expt. Sta. (Cornell Univ.), Mem. 408: 25-42 (mouthparts).
trigonigrnatha Brown. N. C. (Concord). Ecology: Sifted from leaf mold in Berlese funnel.
Am,blyopone (Stigmatom.ma) trigonignatha Brown, 1949. Psyche 56: 81-84. ♂.

Taxonomy: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bui. 122: 169, 185.

Genus PRIONOPELTA Mayr

Prionopelta Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 53: 503.

Type-species: Prionopelta punctulata Mayr. Monotypic.
Renea Donisthorpe, 1947. Ann. and Mag. Nat. Hist. (11) 14: 183. Preocc. by NeviU, 1880.

Type-species: Renea testacea Donisthorpe. Orig. desig.
Examblyopone Donisthorpe, 1949. Ann. and Mag. Nat. Hist. (12) 2: 401.

Type-species: Examblyopone churchilli Donisthorpe. Orig. desig.

A tropicopolitan genus of small, soil-inhabiting ants.

Revision: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bui. 122: 173-178.

Taxonomy: Brown, 1951. Brooklyn Ent. Soc., BuL 46: 102. —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 120 (larvae). —Brown, 1953. Breviora 11: 11. —Wheeler and
Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 447 (larvae; Prirwpelta{!)).

Prionopelta antillana Forel
Fla. (Juniper Springs, Marion Co.); W. Indies, Central America to Brazil,
Bolivia (?). Ecology: Specimens have been foimd in soil.
Prionopelta punctulata antillana Forel, 1909. Deut. Ent. Ztschr., p. 239. ♂.

Taxonomy: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bui. 122: 178.

Biology: Kempf, 1961. Studia Ent. 4: 489-490 (in soil samples in Surinam).

Tribe PLATYTHYREINI
Revision: Brown, 1975. Search, Agr., Ent. (Ithaca) 15, 5 (1): 4-11 (world genera and species).




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Taxonomy: Brown, 1952. Breviora 6: 1-6.

Genus PLATYTHYREA Roger

Platythyrea Roger, 1863. Berlin Ent. Ztschr. 7: 172.

Type-species: Pachycondyla pu7ictata Smith. Desig. by Bingham, 1903.

A tropicopolitan genus with 8 Neotropical species, only one of which reaches the United
States.

Taxonomy: Brown, 1952. Breviora 6: 4 (in key). —Wheeler and Wheeler, 1964. Ent. Soc.
Amer., Ann. 57: 446 (larvae). —Wheeler and Wheeler, 1976. Amer. Ent. Soc, Trans. 102: 41
(revised characterization of larvae).

Platythyrea punctata (Smith)
S. Fla. and s. Tex., s. to W. Indies, Central Amer., and Brazil. Ecology:

Nests in small colonies up to a few hundred individuals each, usually in rotten logs and
stumps or under the bark of trees in shady situations. Workers are active, forage singly,
and are carnivorous and predatory.
Pachycondyla punctata Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 108. 5, cj.
Platythyrea pruinosa Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 962. ♂.

Taxonomy: Forel, 1893. Ent. Soc. London, Trans., p. 358 (worker, female). —Mann, 1916.
Harvard Univ., Mus. Comp. ZooL, Bui. 60: 403. —Wheeler and Wheeler, 1952. Amer.
Midland Nat. 48: 119 (larva).

Biology: Forel, 1899. Rev. Suisse Zool. 9: 335 (in Barbados). —Wheeler, 1905. Amer. Mus.
Nat. Hist., Bui. 21: 80 (in Bahamas). —Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 824
(in Puerto Rico). —Brown, 1957. Harvard Univ., Mus. Comp. Zool., Bui. 116: 229 (in
Mexico).

Tribe ECTATOMMINI
Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 118: 175-362.

Genus ECTATOMMA Smith

Ectatomma Smith, 1859. Cat. Hym. Brit. Mus., v. 6, p. 102.

Type-species: Formica tuberculata Olivier. Desig. by Bingham, 1903.

A Neotropical genus of about 14 species, none of which are native to the United States. One
species was introduced into southern Texas for biological control purposes, but the attempt
failed.

Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 118: 206-211, 295-299.

Taxonomy: Wheeler and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 449 (larvae).

Biology: Weber, 1946. Ent. Soc. Wash., Proc. 48: 1-16 {E. tuberculatum (Olivier) and E.
ruidum (Roger).

Ectatomma tuberculatum (Olivier)
S. Tex. (?); Mexico to n. Argentina. Ecology: Nests are in the soU and
are composed of several hundred individuals. Workers are predatory and carnivorous
and also attend membracids and aphids on plants. Introduced into Victoria Co., Texas
from Guatemala in 1904-1905 to combat the cotton boll weevil. The attempted
introduction led to a fiery confrontation between O. F. Cook, instigator of the
experiment, and W. M. Wheeler. Known as the kelep.

Formica tuberculata Olivier, 1791. Encycl. Meth., Diet. Ins., v. 6, p. 498. ♀.

Formica tridentata Fabricius, 1804. Systema Piezatorum, p. 42. ♂.

Ectatomma fermginetis Norton, 1868. Comm. Essex Institute 6: 5. 5, 6.

Ectatomma tuberculatum var. punctigerum Emery, 1890. Soc. Ent. France, Ann. 10: 56.
5.

Ectatomma tuberculatum var. acrista Forel, 1909. Deut. Ent. Ztschr., p. 254. ♂, ♀.

Ectatomma tuberculatum var. irregularis Santschi, 1921. Soc. Vaud. Nat. des Sci., Bui. 54:
83. ♂.




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Taxonomy: Lever, 1930. Ent. Monthly Mag. 66: 214 (female). —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 129-133 (larva). —Brown, 1958. Harvard Univ., Mus. Comp. Zool.,
Bui. 118: 209, 211, 298-299. — Kempf, 1962. Studia Ent. 5: 2-3.

Biology: Cook, 1904. U. S. Dept. Agr., Div. Ent., Bui. 49: 1-15. —Cook, 1904. Science 19:
862-864. —Cook, 1904. Science 20: 310-312 (pupation). —Wheeler, 1904. Science 20: 437-440
(pupation; feasibility of introduction). —Cook, 1904. Science 20: 611-612. —Wheeler, 1904.
Science 20: 766-768. —Cook, 1905. U. S. Dept. Agr., Bur. Ent., Tech. Ser. 10: 1-55.
—Wheeler, 1905. Science 21: 706-710 (criticism of Cook's work). —Cook, 1906. Science 23:
187-189. —Wheeler, 1906. Science 23: 348-350. —Weber, 1946. Ent. Soc. Wash., Proc. 48:
1-16 (biology and economic significance).

Morphology: Haskins and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37: 100-162 (physiology).
— Gotwald, 1969. N. Y. Agr. Expt. Sta. (Cornell Univ.), Mem. 408: 25-42 (mouthparts).

Genus GNAMPTOGENYS Roger

Gnamptogenys Roger, 1863. Berlin Ent. Ztschr. 7: 174.

Type-species: Ponera tomata Roger. Desig. by Emery, 1911.
Ectatomma subg. Stictoponera Mayr, 1887. Zool.-Bot. (Resell. Wien, Verh. 37: 539.

Type-species: Ectatomma coxale Roger. Desig. by Bingham, 1903.
Ectatomma subg. Holcoponera Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37: 540.

Type-species: Gtvamptogenys striatula Mayr. Desig. by Emery, 1911.
Alfaria Emery, 1896. Soc. Ent. Ital., Bui. 28: 41.

Type-species: Alfaria sim.ulans Emery. Monotypic.
Ectatom,ma subg. Poneracantha Emery, 1897. Mus. Civ. Stor. Nat. (]tenova, Ann. 38: 547.

Type-species: Ectatomma (Holcoponera/) bispinosum Emery. Monotypic.
Rhopalopone Emery, 1897. Mus. Civ. Stor. Nat. Genova, Ann. 38: 549.

Type-sp)ecies: RhopalopoTie epinotalis Emery. Monotypic.
Emeryella Forel, 1901. Soc. Ent. Belg., Ann. 45: 334.

Type-species: Emeryella schmitti Forel. Monotypic.
Ectatomma subg. Mictoponera Forel, 1901. Soc. Ent. Belg., Ann. 45: 372.

Type-species: Ectatomma (Mictoponera) diehli Forel. Monotypic.
Ectatomma subg. Parectatomma Emery, 1911. In Wytsman, Gen. Ins., fasc. 118, p. 44.

Type-species: Ectatomma (Gnamptogenys) triangulare Mayr. Orig. desig.
Spaniopone Wheeler and Mann, 1914. Amer. Mus. Nat. Hist., Bui. 33: 11.

Type-sp)ecies: Spaniopone haytiana Wheeler and Mann. Monotypic.
Wheeleripone Mann, 1919. Harvard Univ., Mus. Comp. Zool., Bui. 63: 282.

Type-species: Wheeleripone albiclava Mann. Orig. desig.
Opisthoscyphus Mann, 1922. U. S. Natl. Mus., Proc. 61: 4.

Type-species: Opisthoscyphus scabrostis Mann. Monotypic.
Ectatxymma subg. Commateta Santschi, 1929. ZooL Anz. 82: 476.

Type-species: Ectatomma (Parectatomma) bruchi Santschi. Orig. desig.
Ectatomma subg. Tammoteca Santschi, 1929. Zool. Anz. 82: 476.

Type-species: Ectatomma concinna Smith. Orig. desig.
Emeryella subg. Barbourella Wheeler, 1930. New England Zool. Club, Proc. 12: 10.

Type-species: Emeryella (Barbourella) banksi Wheeler. Orig. desig.

Brown (1958) treated 81 species of which about 64 are found in the New World tropics and di-
vided the genus into four species groups. The question has been raised as to whether our only
species is native or adventive.

Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 118: 211-241, 299-330.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 133-134 Oarvae). —Wheeler
and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 540 (larvae). —Wheeler and Wheeler, 1976.
Amer. Ent. Soc., Trans. 102: 43 (revised characterization of larvae).

Gnamptogenys hartmani (Wheeler)
La. (Lucky), Tex. (Huntsville); Honduras. Ecology: Specimens have been
taken from soil under banana trees in Honduras and from nests of Trachymyrmex
septentrionalis (McCook) in La.
Ectatomma, (Parectatom.ma) hartmani Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 390.
5.




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Taxonomy: Brown, 1958. Harvard Univ., Mus. Comp. ZooL, Bui. 118: 228, 230, 234, 302.
— Brown, 1961. Psyche 68: 69 (from Honduras; single worker from Tex. described by
Wheeler possibly a locality error or adventive specimen).

Biology: Echols, 1964. Ent. Soc. Amer., Ann. 57: 137 (La.; first reproductive colonies taken in
U. S.).

Genus PROCERATIUM Roger

Proceratium Roger, 1863. Berlin. Ent. Ztschr. 7: 171.

Type-species: Proceratium silaceum Roger. Monotypic.
Sysphingta Roger, 1863. Berlin. Ent. Ztschr. 7: 175.

Type-species: Sysphingta micrommata Roger. Monotypic.
Sysphincta Mayr, 1865. Reise d. Novara, Zool. 1 (1) Formicidae, p. 12. Emend.

About 24 world species are known, most of which are found in the warmer parts of the
northern temperate region of the world. They are hypogaeic, and the small colonies of two or
three dozen individuals are found in well-rotted, moist wood such as that of logs and stumps. A
constant, high moisture content is essential. Toward the cooler areas they may be under the
deepest rocks. Workers are sluggish and carnivorous, apparently feeding almost exclusively on
the eggs of other Arthropods.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 264-266. —Brown, 1958. Harvard
Univ., Mus. Comp. Zool., Bui. 118: 241-248.

Taxonomy: Wheeler aid Wheeler, 1952. Amer. Midland Nat. 48: 135-137. (larvae). —Wheeler
and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 451 (larvae). — Snelling, 1967. Los Angeles
Co. Mus., Contrib. Sci. 124: 1-10 (key to New World species). — Wheeler and Wheeler,
1971. Ent. Soc. Amer., Ann. 64: 1202 (larvae).

Biology: Brown, 1958. Psyche 65: 115 (predation of arthropod eggs).
califomicum Cook. Calif. Found in several scattered localities from Sutter Co. to Los Angeles
Co. The worker is unknown.
Procratiumd) califomicum Cook, 1953. The Ants of Calif., pp. 45-46. ♂.

Taxonomy: Snelling, 1967. Los Angeles Co. Mus., Contrib. Sci. 124: 1-10 (female, male).

Proceratium croceum (Roger)
Va. to Fla. w. to 111., Tex.

Ponera crocea Roger, 1860. Berlin Ent. Ztschr. 4: 288. ♀.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 264 (worker, female). —Smith, 1930.
Ent. Soc. Amer., Ann. 23: 390-392 (male). —Wheeler and Wheeler, 1952. Amer. Midland
Nat. 48: 135 (larva).

Biology: Raskins, 1930. N. Y. Ent. Soc, Jour. 38: 121-126. —Brown, 1958. Harvard Univ.,
Mus. Comp. Zool., Bui. 118: 246-247.

Proceratium melinum (Roger)
"Carolina"; Europe. A European species that doubtfully occurs in North

America and has not been collected here since it was originally described; types may

have been mislabeled.
Ponera melina Roger, 1860. Berlin. Ent. Ztschr. 4: 291. 5, 9, 6.
Sysphingta Europaea Forel, 1886. Soc. Ent. Belg., Ann. (C. R.) 30: CLXIH. 5.
Sysphincta europaea rossica Amoldi, 1930. Zool. Anz. 91: 144. ♂, ♀.
Sysphincta fialai Kratochvil, 1944. In "Mohelno." Arch. Svaz. ochr. prir. dom. Morave,

Svazek 6: 54, 86.

Taxonomy: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 118: 243, 246-248, 334
(doubtful in N. Amer.; also biological notes).

Proceratium pergandei (Emery)
Mass. to Fla. w. to Iowa, Ark., La.

Sysphincta pergandei Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 264. ♂.

Taxonomy: Smith, 1928. Ent. News 39: 242-243 (male). —Cole, 1940. Amer. Midland Nat. 24:
36. —Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 118: 243, 246, 247-248, 336 (also
biological notes).




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Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 276, 304. —Wesson and Wesson, 1940. Amer.
Midland Nat. 24: 90-91 (ate only inside of gaster of dead workers of other ants.)

Proceratium silaceum Roger
Mass., s. Ont. (Pelee Is. and vicinity) s. to n. Fla., w. to 111., Ark., Okla.
Proceratium silaceum Roger, 1863. Berlin. Ent. Ztschr. 7: 172. ♀.
Proceratium crassicome Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 265. ♂.
Procemtium crassicome var. vestitum Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 266. ♂.
Proceratium silaceum rugulosum Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 390. ♂.
9.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 137 (larva). —Brown, 1958.
Harvard Univ., Mus. Comp. Zool., Bui. 118: 241, 245-248, 336 (also biology).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 373, 375. —Wheeler, 1916. Ind. Acad.
Sci., Proc. 26: 460. —Smith, 1928. Ent. News 29: 244. —Dennis, 1938. Ent. Soc. Amer., Ann.
31: 272, 273, 276, 304. —Wesson and Wesson, 1940. Amer. Midland Nat. 24: 91.

Morphology: Kennedy and Talbot, 1939. Ind. Acad. Sci., Proc. 48: 206-210 (each caste; also
biological notes).

Genus DISCOTHYREA Roger

Discothyrea Roger, 1863. Berlin. Ent. Ztschr. 7: 176.

Type-species: Discothyrea testacea Roger. Monotypic.
Pseudosysphincta Arnold, 1916. South. Afr. Mus., Ann. 14: 161.

Type-species: Pseudosysphincta poweri Arnold. Orig. desig.
Prodiscothyrea Wheeler, 1916. Roy. Soc. South. Aust., Trans. 60: 33.

Type-species: Prodiscothyrea velutina Wheeler. Monotypic.
Pseudosphincta Wheeler, 1922. Amer. Mus. Nat. Hist., Bui. 45: 645, 762. Variant spelling of
Pseudosysphincta.

Most of the 26 species of this genus are found in the tropical and southern temperate regions
of the world. Because of their small size and cryptobiotic habits, they are not commonly collected
and their biology is poorly known. The colonies are small and a number of forms have been col-
lected from leaf litter, humus, and rotten logs.

Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 118: 248-253.

Taxonomy: Borgmeier, 1949. Rev. BrasU. Biol. 9: 205. — Borgmeier, 1957. Acad. Brasil. de
Cien., An. 29: 124-125 (male). —Wheeler and Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1202
(larvae).

Biology: Brown, 1958. Psyche 64: 115 (as predators of eggs of other arthropods).

Discothyrea testacea Roger
N. C. to Fla.; Okla. Ecology: Some have been found in pine forest litter and in
soiL
Discothyrea testacea Roger, 1863. Berlin. Ent. Ztschr. 7: 177. ♂, ♀.

Taxonomy: Weber, 1939. Ent. Soc. Amer., Ann. 32: 99 (worker, female). —Smith and Wing,
1954. N. Y. Ent. Soc., Jour. 62: 110-112 (worker, female). —Smith, 1955. Brooklyn Ent.
Soc., Bui. 50: 98 (probable type locality; collector). — Brown, 1958. Harvard Univ., Mus.
Comp. Zool., Bui. 118: 253, 341-342.

Tribe PONERINI

Taxonomy: Brown, 1963. Breviora 190: 1-10.

Genus PACHYCONDYLA Smith

PachycoTidyla Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 105.

Type-species: Formica crassinoda Latreille. Desig. by Emery, 1901.
Neoponera Emery, 1901. Soc. Ent. Belg., Ann. 45: 40, 43.

Type-species: Formica villosa Fabricius. Orig. desig.
Euponera subg. Trachym^sopus Emery, 1911. In Wytsman, Gen. Ins., fasc. 118, p. 84.

Type-species: Formica stigma Fabricius. Orig. desig.




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Brown (1973) has synonymized Neoponera, Trachymesopus, and several other genera with
Pachycondyla. This is a large genus in the tropical regions of the world, but only three species
reach the United States.

Revision: Emery, 1890. Soc. Ent. France, Ann. 10: 71-74 (Pachycondyla).

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 119: 352 (Trachymesopus
not a subgenus of Euponera). — Kempf, 1960. Studia Ent. 3: 423-428 (New World species
of Trachymesopus). —Kempf, 1961. Rev. Brasil. Ent. 10: 89-204 (Brazilian species of
Pachycondyla). — Brown, 1963. Breviora 190: 6-8 (three species groups of
Trachymesopus). —Kempf, 1964. Studia Ent. 7: 49-52 (key to species of Pachycondyla).
— Brown, 1973. In Meggers, et al.. Tropical forest ecosystems in Africa and S. Amer., pp.
178-185 (generic synonymy).

Pachycondyla harpax (Fabricius)
La., Tex. s. to Brazil; W. Indies. Ecology: Colonies of about 150 individuals
are found in rotten logs and stumps or in soil beneath objects. Workers avoid direct
sunlight and forage in the morning and in shade for other insects and myriapods on
which they feed. There are both ergatoid and normal females.

Formica harpax Fabricius, 1804. Systema Piezatorum, p. 401. ♂.

Pachycondyla Montezumia Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 108. ♀, ♂.

Pomerad) amplinoda Buckley, 1866. Ent. Soc. Phila., Proc. 6: 171. ♂.

Pachycondyla Orizabana Norton, 1868. Amer. Nat. 2: 64. ♀.

Pachycondyla harpax var. dibullana Forel, 1901. Rev. Suisse Zool. 9: 347. 2 .

Pachycondyla harpax var. irina Wheeler, 1925. Arkiv for Zool. 17A: 5. ♀.

Pachycondyla harpax var. concinna Wheeler, 1925. Arkiv for Zool. 17 A: 5. ♂, ♀.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 401-403 (each caste). —Brown,
1950. Wasmann Jour. Biol. 8: 247-248 (species synonymy). —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 617-618 (larva). —Kempf, 1961. Rev. Brasil. Ent. 10: 194.

Biology: Wheeler, 1900. Biol. Bui. 2: 1-31. — Haskins and Enzmann, 1938. N. Y. Acad. Sci.,

Ann. 37: 150-151 (formation of new colony; in Canal Zone).

Pachycondyla stigma (Fabricius)
Fla.; W. Indies, Mexico s. to n. Argentina; S. China to Samoa, n.

Queensland. Ecology: They prefer to nest in moist, dead logs or stumps, occasionally
under stones. Apparently a tramp species distributed by commerce outside the New
World. Whether or not it is endemic or adventive to Florida is unknown.

Formica stigma Fabricius, 1804. Systema Piezatorum, p. 400. ♀.

Ponera quadridentata. Smith, 1859. Linn. Soc. London, Jour. Zool. 3: 143. ♀.

Ponera Americana Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 13: 722. ♂.

Euponera (Trachymesopus) nixoni Donisthorpe, 1943. Ann. and Mag. Nat. Hist., ser. 11,
10:441. ♀.

Euponera (Trachymesopus) brunneus Donisthorpe, 1947. Ann and Mag. Nat. Hist., ser. 11,
14: 300-301. ♂, ♀.

Euponera (Trachymesopus) sexdentatus Donisthorpe, 1949. Ann. and Mag. Nat. Hist., ser.
12, 1: 746. ♀.

Taxonomy: Smith, 1934. Ent. Soc. Amer., Ann. 27: 561-564. —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 627 (larva). —Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bui.
119: 355 (synonymy and distribution). — Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 22
(Polynesia). — Wheeler and Wheeler, 1976. Amer. Ent. Soc, Trans. 102: 55-58 (descriptions
of larvae from different localities; as Mesoponera stigma).

Biology: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 824. —Haskins and Enzmann, 1938.

N. Y. Acad. Sci., Ann. 37: 151 (colony formation; Canal Zone). —Kempf, 1960. Studia Ent.

3: 427-428. — WUson, 1959. Evolution 13: 128 (distribution; ecology).

Pachycondyla villosa (Fabricius)
S. Tex. (as far north as San Antonio) s. to n. Argentina. Ecology: Colonies
occur in the soil and in logs and stumps. Workers run rapidly in the bright sun in search
of insects on which they feed. They can sting severely. The largest ponerine ant in the
U.S.

Formica villosa Fabricius, 1804. Systema Piezatorum, p. 409. ♀.

Ponera bicolor Guerin, 1845. Iconogr. Regne Anim., Ins., v. 7, p. 242. ♀.

Ponera pilosa Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 95. ♂.




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Ponera pedunculata Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 96. ♂.

Taxonomy: Roger, 1861. Berlin Ent. Ztschr. 5: 1 (worker, male). —Wheeler and Wheeler,
1952. Amer. Midland Nat. 48: 615 (larva). —Wheeler and Wheeler, 1971. Ent. Soc. Amer.,
Ann. 64: 1205 (larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 404.

Genus BRACHYPONERA Emery

Euponera subg. Brachyponera Emery, 1901. Soc. Ent. Belg., Ann. 45: 43.
Type-species: Ponera sennaarensis Mayr. Orig. desig.

A small genus found in the Old World. The one species in North America is adventive.

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 119: 346 (not a subgenus of
Euponera). — Brown, 1958. Acta Hym. 1: 21.

Brachyponera solitaria (Smith)
Va., N. C, Ga.; Japan, China, and adjacent areas. Ecology: The small colonies
are found in moist, rotten wood or in soil beneath objects; they prefer dark, damp
places. Food consists of small arthropods. Accidentally introduced into N. Amer.
Ponera solitaria Smith, 1874. Ent. Soc. London, Trans., p. 404. ♂.

Taxonomy: Smith, 1934. Ent. Soc. Amer., Ann. 27: 558-561 (description; first record for N.
Amer.). —Wheeler and Wheeler, 1952. Amer. Midland Nat. 49: 629 (larva). —Wheeler and
Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1207 (larva).

Biology: Smith, 1934. Ent. Soc. Amer., Ann. 27: 560-561.

Genus CRYPTOPONE Emery

Cryptopone Emery, 1892. Soc. Ent. France, Bui. 61: CCLXXV.

Type-species: Amblyopone^ testacea Motschulsky. Monotypic.

Most species of this genus are found in Southeast Asia and vicinity; only one, apparently en-
demic species occurs in the United States.

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool, Bui. 119: 357-361. —Brown, 1963.
Breviora 190: 6.

Cryptopone gilva (Roger)
Ga., Tenn., Ala., Miss., Ark., La., Tex. Ecology: The smaU colonies nest in moist
dead logs or stumps, preferring loose frass under bark.
Ponera gilva Roger, 1863. Berlin. Ent. Ztschr. 7: 170. ♂.
Euponera (Trachymesopus) gilva hamedi Smith, 1929. Ent. Soc. Amer., Ann. 22: 543. ♂.

Taxonomy: Smith, 1934. Ent. Soc. Amer., Ann. 27: 561-563. — Creighton and Tulloch, 1930.
Psyche 37: 73-79 (each caste). —Wheeler and Wheeler, 1952. Amer. Midland Nat. 48:
625-627 Garva). —Brown, 1963. Breviora 190: 6.

Biology: Haskins, 1931. N. Y. Ent. Soc., Jour. 39: 507-521. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31: 277, 304.

Genus PONERA Latreille

Ponera Latreille, 1804. Nouv. Diet. Hist. Nat. 24: 179.

Type-species: Formica contractu Latreille. Desig. by Latreille, 1805.
Pseudocryptopone Wheeler, 1933. Amer. Mus. Novitates 672: 12-13.

Type-sf)ecies: Cryptopone tenuis Emery. Orig. desig.
Selenopane Wheeler, 1933. Amer. Mus. Novitates 672: 19.

Type-species: Ponera selenophora Emery. Orig. desig.

Taylor (1967) treated 28 world species and divided the genus into several species groups. Most
species are found in the Indo-Australian area The only two New World species are North
American. Most forms are found in forested areas where they nest in small colonies in rotten
wood or stumps or in the soil beneath cover. The workers are carnivorous.

Revision: Smith, 1936. Ent. Soc. Amer., Ann. 29: 420-430 (in part; U. S.). —Taylor, 1967.
Pacific Ins. Monog. 13: 1-112 (world).




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Taxonomy: Wilson, 1957. Harvard Univ., Mus. Comp. Zool., Bui. 116: 356-357. —Brown, 1958.
Acta Hym. 1: 22-23. —Wheeler and Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1208 (larvae).

Species Group Leae

Ponera exotica Smith
N. C, Okla. Ecology: Specimens are all from Berlese samples of leaf litter or
leaf mold. Possibly introduced. Affinities are with the I ndo- Australian fauna.
Ponera exotica Smith, 1962. Acta Hym. 1: 378-382. ♂, ♀.

Taxonomy: Taylor, 1967. Pacific Ins. Monog. 13: 96-97.

Species Group Coarctata

Ponera pennsylvanica Buckley
N. S., Que. s. to Fla. w. to Ont., N. Dak., Colo., Utah, N. Mex. Ecology:
Most abimdant in the eastern deciduous forests, east of the 97th meridian, with only
scattered records in the western states. Nests are found under rotting logs, in rotting
stumps, small fragments of wood, acorns and other objects, or in soil or leaf mold. In
drier habitats they may nest under stones. Workers forage in or on the ground and are
carnivorous. Only occasionally a household pest.
Ponera Pennsylvanica Buckley, 1866. Ent. Soc. Phila., Proc. 6: 171. ♀.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 631 Garva). —Taylor, 1967.
Pacific Ins. Monog. 13: 29-38 (also biology, ecology). — Wheeler and Wheeler, 1971. Ent.
Soc. Amer., Ann. 64: 1210 (larva).

Biology: Wheeler, 1900. Biol. Bui. 2: 22-23, 43-56. —Wheeler, 1917. Conn. State Geol. and Nat.
Hist. Survey Bui. 22: 581. — HasWns and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37: 152-155
(colony formation). — Headley, 1952. Ent. Soc. Amer., Ann. 45: 436-438. — Kannowski, 1959.
Insectes Sociaux 6: 118 (colony founding). — Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 91-92. —Gregg, 1963. Ants of Colo., pp. 281-282, 284.

Morphology: Gotwald, 1969. N. Y. Agr. Expt. Sta. (Cornell Univ.) Mem. 408: 25-42
(mouthparts).

Genus HYPOPONERA Santschi

Ponera subg. Hypoponera Santschi, 1938. Soc. Ent. France, Bui. 43: 79.
Type-species: Ponera aheillei Andre. Orig. desig.

A large cosmopolitan genus whose habits are similar to those of species of Ponera. Prior to
Taylor, 1967, the species below were assigned to the genus Ponera.

Revision: Smith, 1936. Ent. Soc. Amer., Ann. 29: 420-430 (in part, as Ponera; U. S.).

Taxonomy: Taylor, 1967. Pacific Ins. Monog. 13: 9-14 (a distinct genus). —Taylor, 1968. Ent.
News 79: 63-66 (list of N. Amer. species). —Wheeler and Wheeler, 1971. Ent. Soc. Amer.,
Ann. 64: 1210 Garvae).

Hypoponera gleadowi (Forel)
Md. (Priest Bridge); Asia. Apparently a widespread tramp species. Possibly
introduced. No other confirmed records from the U. S.
Ponera Gleadowi Forel, 1895. R. Accad. Sci. Bologna, Mem. 5: 292-293. ♂.
Ponera japonica formosae Forel, 1913. Arch. f. Naturgesch. 79: 186.
Ponera oblongiceps Smith, 1939. Ent. Soc. Wash., Proc. 41: 76-78. 9,9, apterous ergatoid
6.

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bui. 119: 326 (Melanesia;
provisional synonymy). —Taylor, 1967. Pacific Ins. Monog. 13: 11, 12, 76. —Wilson and
Taylor, 1967. Pacific Ins. Monog. 14: 29 (unknown from Polynesia; synonymy given for

Hypoponera gleadowi
***authority mismatch
by Wilson, 1958 provisionally transferred to synonymy of punctatissinw,
(Roger)). —Taylor, 1968. Ent. News 79: 65 (U. S.).

Hypoponera inexorata (Wheeler)
S. C. to Fla. w. to Tex., Ariz.; s. to Central Amer.
Ponera inexorata Wheeler, 1903. Psyche 10: 94. ♂, ♀.

Taxonomy: Taylor, 1968. Ent. News 79: 65.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 406.




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Hypoponera opaciceps (Mayr)
S. C. to Fla. w. to Colo., Ariz.; s. to Argentina, W. Indies; s. e. Asia,
Polynesia. Probably spread from New World to Old World by commerce.
Ponera opaciceps Mayr, 1887. Zool.-Bot. Gesell. Wien., Verb. 37: 536. ♀.
Ponera perkinsi Forel, 1899. Faiina Hawaiiensis, p. 117. 5, 9, d.
Ponera andrei Emery, 1900. Termes. Fuzetek 23: 318. ♂.

Taxonomy: Smith, 1929. Ent. Soc. Amer., Ann. 22: 545-546. —Smith and Haug, 1931. Ent.
Soc. Amer., Ann. 24: 507-509 (ergatandrous form). — Wheeler and Wheeler, 1952. Amer.
Midland Nat. 48: 364 (larva). — Kempf, 1962. Studia Ent. 5: 7-9. —Wilson and Taylor, 1967.
Pacific Ins. Monog. 14: 28 (Polynesia). —Taylor, 1968. Ent. News 79: 65. —Wheeler and
Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1210 (larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 125, 404. —Smith, 1927. Ent. News
38: 308-309. —Kempf, 1960. Studia Ent. 3: 391. —Gregg, 1963. Ants of Colo., pp. 283-284.

Hypoponera opacior (Forel)
Va. to Fla. w. to Ohio, Iowa, Colo., Tex.; Oreg., Calif.; Mexico s. to Chile,

Argentina; W. Indies. More sporadically distributed west of Tex. than in eastern states.
Ponera trigona var. opacior Forel, 1893. Ent. Soc. London, Trans., p. 363. ♂, ♀.

Taxonomy: Taylor, 1968. Ent. News 79: 65.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 272, 274, 277, 304 (Tenn.) —Cole, 1940.
Amer. Midland Nat. 24: 37 (Smoky Mts.). — Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 63
(Calif.). —Potts, 1948. Pan-Pacific Ent. 24: 26 (Calif.). —Gregg, 1963. Ants of Colo., pp.
284-286.

Hypoponera punctatissima (Roger)
Fla., Tex., N. Mex., Ariz., Calif.; s. to Central Amer., W. Indies;

Europe, N. Africa. Ecology: Nearly cosmopolitan in warmer parts of world. Possibly

introduced; probably of African origin.
Ponera punctatissima Roger, 1859. Berlin. Ent. Ztschr. 3: 246-248. $ , 9 .
Ponera androgyna Roger, 1859. Berlin. Ent. Ztschr. 3: 254. Ergatoid 6.
Ponera ergatandria Forel, 1893. Ent. Soc. London, Trans., p. 365. 5,9, apterous ergatoid

6.
Ponera punctatissima schauinslandi Emery, 1899. Zool. Jahrb., Abt. f. System 12: 439. ♀.

Taxonomy: Smith, 1936. Ent. Soc. Amer., Ann. 29: 422, 425-426 (female, worker, apterous
ergatoid male). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 28-29 (Polynesia;
synonymy under gleadovn (Forel) listed by Wilson, 1958, Harvard Univ., Mus. Comp. Zool.,
Bui. 119: 328-329 provisionaUy transferred to punctatissima. This synonymy is not listed
above). —Taylor, 1968. Ent. News 79: 65.

Biology: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 825. —Mallis, 1941. South. Calif.
Acad. Sci., Bui. 40: 63.

Genus LEPTOGENYS Roger

Only a single species of this tropicopolitan genus is known to occur in the United States.

Taxonomy: Wheeler and Wheeler, 1976. Amer. Ent. Soc., Trans. 102: 49 (revised
characterization of larvae).

Genus LEPTOGENYS Subgenus LEPTOGENYS Roger

Leptogenys Roger, 1861. Berlin. Ent. Ztschr. 5: 41.

Type-species: Leptogenys falcigera Roger. Desig. by Bingham, 1903.
Dorylozelus Forel, 1915. Arkiv for Zool. 9: 24-25.

Type-species: Dorylozelus joebergi Forel. Monotypic.
Microbolbos Donisthorpe, 1948. Entomologist 81: 170.

Type-species: Microbolbos testacev^ Donisthorpe. Orig. desig.

Not known to occur in the Nearctic Region.

Genus LEPTOGENYS Subgenus LOBOPELTA Mayr

Lobopelta Mayr, 1862. Zool.-Bot. GeseU. Wien, Verb. 12: 733.

Type-species: Ponera diminuta Smith. Desig. by Bingham, 1903.




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Revision: Wheeler, 1923. Amer. Mus. Novitates 90: 1-16.

Leptogenys elongata
***authority mismatch
elongata (Buckley). La., Tex. Ecology: The small colonies are found in the soil. The
workers apparently forage singly and feed largely, if not exclusively, on pillbugs. There
is no typical female as with most ants, reproduction being carried on by a wingless,
slightly modified worker form with an enlarged gaster.
Ponera Texana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 170. $ . A questionable synonym.
Ponera elongata Buckley, 1866. Ent. Soc. Phila., Proc. 6: 172. ♂.
Lobopelta septentrionalis Mayr, 1866. Zool.-Bot. Gesell. Wien, Verb. 36: 438. ♂.

Taxonomy: Wheeler, 1904. Biol. Bui. 6: 257-259 (worker, gynaecoid female, male). —Wheeler
and Wheeler, 1952. Amer. Midland Nat. 48: 641 (larva). —Wheeler, 1900. Biol. Bui. 2: 1-31.

Morphology: Hermann, 1969. Kans. Ent. Soc., Jour. 42: 239-243 (poison apparatus).

Leptogenys elongata
***authority mismatch
manni Wheeler. Fla.

Leptogenys {Lobopelta) elongata manni Wheeler, 1923. Amer. Mus. Novitates 90: 14-15. ♂.

Tribe ODONTOMACHINI

Genus ODONTOMACHUS Latreille

Odontomachus Latreille, 1804. Nouv. Diet. Hist. Nat. 24: 179.
Type-sp)ecies: Formica haematoda Linnaeus. Monotypic.

Four forms of this tropicopolitan genus reach the southern portions of the United States. All
of these have previously been considered as subspecies of the Neotropical O. haematodus (L.).
Most colonies are small and are found in soil or in rotting logs and stumps. Workers are
predaceous and carnivorous. Species of this genus have elongated, linear mandibles and long
hairs which arise between the bases of the mandibles and point forward. These hairs act as trig-
gers when the mandibles are open. When the hairs are touched, the mandibles snap shut result-
ing in a clicking sound. If the mandibles close suddenly on a small object, the object may be cut
in two; if they close suddenly on a large object and the mandibles slide over it, the ant is thrown
in a series of leaps by the force of its closing mandibles.

Revision: Smith, 1939. N. Y. Ent. Soc., Jour. 47: 125-130 (U. S.).

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 646 (larvae). —Wheeler and
Wheeler, 1964. Ent. Soc. Amfer., Ann. 57: 455-456 flarvae). —Wheeler and Wheeler, 1971.
Ent. Soc. Amer., Ann. 64: 1212 (larvae). —Wheeler and Wheeler, 1976. Amer. Ent. Soc.,
Trans. 102: 61 (revised characterization of larvae).

Biology: Wheeler, 1900. Biol. Bui. 2: 1-31. —Wheeler, 1922. Amer. Mus. Nat. Hist., Bui. 45:
99-103. — Weyer, 1930. Zool. Anz. 90: 49-55 (leaping habits).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bui. 116: 475-476
(proventriculus).
clams Roger. La., Tex.; Mexico, Clarion Is., W. Indies. Ecology: Found in semi-desert regions
where colonies occur in coarse, gravelly soil, fully exposed to the sun.
Odontomachus clams Roger, 1861. Berlin. Ent. Ztschr. 5: 26. ♂.
Odontomachus texana Buckley, 1867. Ent. Soc. Phila., Proc. 6: 355. ♂.
Odontomachus haematoda clarionensis Wheeler, 1934. Pan-Pacific Ent. 10: 141. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 407 (worker, female, male).
—Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 648-650 (larva). —Taylor and
Wilson, 1961. Psyche 68: 142.

Biology: Wheeler, 1900. Biol. Bui. 2: 1-31. — Haskins and Enzmann, 1938. N. Y. Acad. Sci.,
Ann. 37: 100-143.

Odontomachus coninodis Wheeler
Ariz. (Huachuca Mtns.). Ecology: Small colonies are in coarse gravelly soil
under stones at high elevations, usually over 5000 ft.
Odontomachus haematoda coninodis Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 391.




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Odontomachus desertorum Wheeler
N. Mex., Ariz.; Mexico. Ecology: Small colonies are in coarse gravelly soil
under stones. Found at lower elevations than coninodis.
Odontomachus kaematoda desertorum Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 391.

5.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27: 394.

Odontomachus insularis Guerin
Ga., Fla., Ala.; Mexico to Brazil, W. Indies, Cocos Is., Clipperton Is. Ecology:
Colonies may be in soil or in rotting logs and stumps. Possibly adventive in the U. S.,
distributed by commerce.
Odontomachus insularis Guerin, 1844. Iconogr. Regne Anim. Ins., v. 7, p. 423. ♂.

Taxonomy: Roger, 1861. Berlin. Ent. Ztschr. 5: 26 (worker, male). —Taylor and Wilson, 1961.
Psyche 68: 142 (distribution).

Biology: Haskins and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37: 149 (formation of new
colonies).

Subfamily PSEUDOMYRMECINAE

A small subfamily with one genus in the New World and several genera in the Old World
tropics. M. R. Smith (1951) named this subfamily Leptaleinae based on the genus Leptalea
Erichson, but later (1952) found an earlier valid generic name, Pseudomyrmex Lund, and
changed the subfamily name to Pseudomyrmecinae which has become vddely established.

Taxonomy: Smith, 1951. U. S. Dept. Agr., Agr. Monog. 2: 788 (Leptaleinae). —Smith, 1952.
Ent. Soc. Wash., Proc. 54: 97-98 (Pseudomyrmecinae). —Wheeler and Wheeler, 1973.
Psyche 80: 204-211 (larvae).

Genus PSEUDOMYRMEX Lund

Pseudomyrme Lund, 1831 (June). Ann. Sci. Nat. Zool. 23: 137. Latreille ms.; vernacular.
Pseudom.yrmex Lund, 1831 (November). Notizen aus dem (Jebiete der Natur und
Heilkunde 32(7): 106. Latreille ms.

Type-species: Formica gracilis Fabricius. Desig. by Smith, 1952.
Leptalea Erichson, 1839. Arch. f. Naturgesch. 5: 309. Klug ms.

Type-species: Formica gracilis Fabricius. Desig. by Wheeler, 1911.
Myrmex Guerin, 1844. Iconogr. Regne Anim., Ins., v. 7, p. 427. Preocc. by Sturm, 1826.

Type-species: Formica (Myrmex) perboscii Guerin. Monotypic.
Pseiuiom,yrmxi Guerin, 1844. Iconogr. Regne Anim., Ins., v. 7, p. 427. emend.
Leptalaea Spinola, 1851. Accad. Sci. Torino, Mem. 13: 68. Emend.

Over 190 forms of Psevxkrmyrmex have been described, but only five reach the southern por-
tions of the United States. These ants prefer to nest almost exclusively in preformed plant cavi-
ties such as twigs and branches of trees, stems of plants, acacia thorns, and in culms of sedges
and grasses. Some species may be restricted to one species of plant or even to one part of the
plant. Among these are the acacia nesting species which nest in the swollen bases of the larger
spines. Janzen (1967) studied a case of obligatory mutualism between a species of Pseudomyr-
mex and Acacia in Mexico and fotind that the ant is dependent on the acacia for food and the
acacia is dependent on the ant for a substantial part of its life in order to produce seeds and
become part of the reproductive population. The ant colony protects the plant from damaging
factors such as defoliators and thereby assures its own survival on the plant which is its only
food source.

Revision: Mayr, 1870. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 61: 40&413.

Taxonomy: Smith, 1952. Ent. Soc. Wash., Proc. 54: 97-98 (correct generic name). — Creighton,

1955. N. Y. Ent. Soc, Jour. 63: 19-20 (key to workers of U. S.). —Wheeler and Wheeler,

1956. Ent. Soc. Amer., Ann. 49: 376-379 (larvae). — Kempf, 1958. Studia Ent. (n. s.) 1: 434
(gracilis (Fabricius) group).

Biology: Wheeler, 1913. 2nd Intematl. Cong. Ent., Oxford, Trans. 2: 109-139 (observations on
Central American Acacia ants). — Wheeler and Bailey, 1920. Amer. Phil. Soc., Trans, (n. s.)
22: 235-279 (feeding habits). —Brown, 1960. Ecology 41: 589-592 (ants, acacias and




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browsing mammals), ^anzen, 1966. Evolution 20: 249-275 (coevolution of mutualism
between ants and acacias in Central America). — Janzen, 1967. Kans. Univ., Sci. Bui. 48:
315-558 (interaction of the bull's-hom acacia with an ant inhabitant {Pseudamyrmex
ferruginea Smith) in eastern Mexico). Janzen, 1967. Ecology 48: 26-35 (fire, vegetation
structure and the ant acacia interaction in Central America). — Janzen, 1969. The Condor
71: 240-256 (the ant acacia interaction and birds in Central America.).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bui. 116: 452-453

(proventriculus). — Gotwald, 1969. N. Y. Agr. Expt. Sta (Cornell Univ.), Mem. 408: 128
(mouthparts).

Pseudomyrmex apache Creighton
S. Tex., s. N. Mex., s. Ariz., s. Calif.; n. Mexico. Ecology: Most colonies have
been found in sizeable limbs or trunks of species of Quercus and Prosopis.
Psevdomyrmex apache Creighton, 1952. Psyche 59: 134-139. 5, 9, 6.

Taxonomy: Wheeler and Wheeler, 1956. Ent. Soc. Amer., Ann. 49: 380 (larva).

Biology: Creighton, 1954. Psyche 61: 9-15. —Creighton, 1963. Amer. Mus. Novitates 2156: 1-4
(high mortality during cold snap).

Pseudomyrmex brunneus (Smith)
N. C. to Fla. w. to Tex. s. to Central Amer.; W. Indies. Ecology: Colonies
have been found in dead twigs of various plants and in culms of grasses and sedges.
Pseudomyrma brunnea Smith, 1877. Ent. Soc. London, Trans., p. 63. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 420-421. —Wheeler and Wheeler,
1956. Ent. Soc. Amer., Ann. 49: 382 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 69. —Wheeler, 1932. N. Y. Ent.
Soc., Jour. 40: 3.

Pseudomyrmex elongatus (Mayr)
Fla., Tex. s. to S. Amer.; W. Indies. Ecology: Colonies have been found in
twigs of trees and in culms of grasses and sedges.
Psevdomyrma elongata Mayr, 1870. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 61: 408,

413. ♂.
Pseupomyrma elongata var. cubaensis Forel, 1901. Soc. Ent. Belg., Ann. 45: 342 9 .
Pseudomyrma elongata var. tandem, Forel, 1906. Soc. Ent. Belg., Ann. 50: 228. ♀.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 85-87 (each caste). —Creighton,
1955. N. Y. Ent. Soc., Jour. 63: 17-20. —Wheeler and Wheeler, 1956. Ent. Soc. Amer., Ann.
49: 384 Oarva).

Biology: Wheeler, 1932. N. Y. Ent. See., Jour. 40: 4.
grracilis mexicanus (Roger). Fla., Tex. s. to Central Amer. Ecology: Colonies have been found
in dead limbs of live oaks, in live oak twig galls, in shrubs, hollow stems of composites,
and cavities in other plants. Probably accidentally introduced into Florida. P. gracilis
gracilis (Fabricius) is found in Central and S. Amer.
Psetidomyrma mexicana Roger, 1863. Berlin. Ent. Ztschr. 7: 178. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 421 (worker, female). —Wheeler
and Wheeler, 1956. Ent. Soc. Amer., Ann. 49: 385-386 (larva). —Brown, 1957. Harvard
Univ., Mus. Comp. Zool., Bui. 116: 235.

Biology: Wheeler, 1901. Soc. Ent. Belg., Ann. 45: 204. —Mitchell and Pierce, 1912. Ent. Soc.

Wash., Proc. 14: 69. — Whitcomb, Denmark, Buren, and Carroll, 1972. Fla. Ent. 55: 31-33

(in Florida).

Pseudomyrmex pallidus (Smith)
N. C. to Fla. w. to Ariz., Calif, s. to S. Amer.; W. Indies. Ecology: Colonies
have been foimd in twigs and branches of various plants and in culms of grasses and
sedges. Some of the early western records of this species may actually refer to apache
Creighton. References to P.flavidulus (Smith) for the U. S. pertain to this species;
flavidulus was described from S. Amer. and its correct status is not known (see
Creighton, 1950).

Psevdomyrma pallida Smith, 1855. Ent. Soc. London, Trans., 3: 160. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 419-420. —Wheeler, 1905. Amer.
Mus. Nat. Hist., Bui. 21: 83-85 (each caste). —Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bui. 104: 80-82. —Wheeler and Wheeler, 1956. Ent. Soc. Amer., Ann. 49: 386 (larva).




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Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 69. —Wheeler, 1932. N. Y. Ent.
Soc, Jour. 40: 4.

Morphology: Blum and Callahan, 1963. Psyche 70: 69-74 (morphology and physiology of
poison glands and venom).

Unplaced Taxon of Pseudomyrmecinae
Ponera (EctatomaU)) Lincecumii Buckley, 1866. Ent. Soc. Phila., Proc. 6: 172. 5 . Cent. Tex.

Subfamily MYRMICINAE

This is the largest subfamily of ants and is found throughout the world. In North America, the
Myrmicinae are better represented in the central and southern United States with their in-
cidence rapidly decreasing northwards where the Formicinae become the dominant subfamily.
Members of this subfamily are recognized by the two-segmented petiole and the frontal carinae
which are distant from each other and each of which usually bears a lobe concealing the antennal
insertions.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Amer., Ann. 53:98-110 (larvae). —Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:1-32 (larvae). — Ettershank, 1966. Austral.
Jour. Zool. 14:73-171 (generic revision of world Myrmicinae related to Solenopsis and
Pheidologeton). —Wheeler and Wheeler, 1973. Psyche 80: 204-211 (supp. studies on
larvae). —Wheeler and Wheeler, 1973. Psyche 80: 70-82 (larvae of four tribes, 2nd supp.;
Leptothoracini, Ocymyrmecini, Tetramoriini, Cryptocerini). — Wheeler and Wheeler, 1973.
Ga. Ent. Soc., Jour. 8: 27-39 (larvae of six tribes, 2nd supp.).

Morphology: Blum, 1974. N. Y. Ent. Soc., Jour. 82: 141-147 (Myrmicine trail pheromones:
specificity, source, and significance). — McCluskey, 1974. N. Y. Ent. Soc, Jour. 82: 93-102
(generic diversity in phase of rhythm in Myrmicine ants).

Tribe MYRMICINI

Genus MYRMICA Latreille

Myrmica Latreille, 1804. Nouv. Diet. Hist. Nat. 24:179.

Type-species: Formica rubra Linnaeus. Desig. by Latreille, 1810.

This holarctic genus is found as far north as Labrador and Alaska in North America and is
restricted to higher elevations in the southern parts of its range. There are apparently no sub-
tropical or xerophilous representatives. The moderate sized colonies nest in soil, rotten wood, or
under cover of various objects. Workers are carnivorous but also feed on honeydew of Homop-
tera and exudates of plants. Workers of some species differ from each other very slightly and
males are sometimes needed for determination. Some species are closely related to Palearctic
forms and have been regarded as subspecies of them by some authors.

Revision: Weber, 1947. Ent. Soc. Amer., Ann. 40:437-474. —Weber, 1948. Ent. Soc. Amer.,
Ann. 41:267-308. —Weber, 1950. Ent. Soc. Amer., Ann. 43:189-226.

Taxonomy: Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. 5:73-83 (varieties of M. brevinodis).
—Wheeler and Wheeler, 1952. Psyche 59:112-123 (larvae). —Yarrow, 1955. Roy. Ent. Soc.
London, Proc., Ser. B: Taxonomy 24:113-115 (type-species). — Collingwood, 1958. Roy Ent.
Soc. London, Proc., Ser. A 33:65-75 (Britain). —Wheeler and Wheeler, 1963. Ants of N.
Dak., pp. 94-108. —Collingwood, 1974. Soc. Brit. Ent., Trans. 16:96-101 (Britain).

Biology: Brian, 1957. Insectes Sociaux 4:177-190 (growth and development of colonies).
—Weir, 1958. Insectes Sociaux 5:97-128, 316-339 (polyethism in workers). —Weir, 1958.
Jour. Ins. Physiol. 1:352-360 (effect of temperature variation on queen oviposition and
colony formation). — Kannowski, 1959. Insectes Sociaux 6:143-144 (flight activities and
colony founding). — Weir, 1959. Insectes Sociaux 6:271-290 (influence of worker age on
trophogenic larval dormancy). — Weir, 1959. Insectes Sociaux 6:167-201 (egg masses and
early larval growth). — Weir, 1959. Physiol. Zool. 32:63-77 (interrelation of queen and
worker oviposition). — Carr, 1962. Insectes Sociaux 9:177-211 (influence of queen). — Brian
and Hibble, 1963. Insectes Sociaux 10:71-82 (larval size and influence of queen on growth).




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— Kannowski, 1970. Ent. Soc. Amer., N. Central Branch, Proc. 25:119-125 (colony
populations of 5 species).

Morphology: Weir, 1957. Quart. Jour. Micros. Sci. 98:499-506 (functional anatomy of mid-gut
of larvae). — Weir, 1959. Insectes Sociaux 6:375-386 (changes in the rectro-cerebral
endocrine system of larvae and their relation to larval growth and development). — Crewe
and Blum, 1970. Ztschr. f. Vergleich. Physiol. 70:363-373 (alarm pheromones of 9 species).

Myrmica americana Weber
Que., Maine s. to N. C, Tenn., w. to Man., Colo., Utah, Ariz. Ecology: More

common on east slop)es of Rocky Mtns.; records from west of the Rockies are rare.

Commonly found in grasslands where nests are in soil in open or under objects; food is

varied, consisting of animal matter and plant juices. Sometimes considered as a

subspecies of the Palearctic Myrmica sabuleti Meinert.
Myrmica sabuleti americana Weber, 1939. Lloydia 2:144. 9, 9, S.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:116 (larva).

Biology: "Wheeler, 1916. Conn. State Geol. and Nat. Hist. Survey Bui. 22:587. — Buren, 1944.
Iowa State Col., Jour. Sci. 18:282-283. —Kannowski, 1956. Amer. Midland Nat. 56:175.
— Kannowski and Kannowski, 1957. Ohio Jour. Sci. 57:371-374 (mating activities).
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 96-99. —Gregg, 1963. Ants of Colo., pp.
308-310. — Ayre, 1963. Canad. Ent. 95:712-715 (feeding habits). —Bums, 1964. Ent. Soc.
Amer., Ann. 57:138 (association with tuliptree scale). — Ayre, 1968. Canad. Ent. 100:1(55-172
(prey finding, capture, and transport). — Ayre, 1969. Canad. Ent. 101:118-128 (trail
formation and group foraging). — Ayre, 1971. Ztschr. f. Angew. Ent. 68:295-299 (foraging
and nesting habits).

Myrmica brevispinosa
***authority mismatch
brevispinosa Wheeler. N. Dak., Nebr. s. to Colo., N. Mex., w. to Alta., Idaho.

Ecology: Prefers to nest in stream valleys or on shores of permanent bodies of water.
Myrmica rubra brevinodis var. brevispinosa Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. 5:74.

Myrmica rubra brevinodis var. decedens Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. 5:75. ♂.

S.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:117 (larva). — Kannowski, 1956. Amer.
Midland Nat. 56:176. —Gregg, 1961. N. Y. Ent. Soc., Jour. 69:211.

Biology: Weber, 1942. Canad. Ent. 74:62. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart.
11:241. —Gregg, 1963. Ants of Colo., pp. 296-297, 299. —Wheeler and Wheeler, 1963. Ants
of N. Dak., pp. 101-103 (subspecies not recognized).

Myrmica brevispinosa
***authority mismatch
discontinua Weber. Newfoundland, N. S. w. to mtns. of Wyo., Colo., N. Mex.
Ecology: Apparently more tolerant than the typical subspecies of higher altitudes in
Rockies where the two subspecies overlap.
Myrmica brevinodis discontinue Weber, 1939. Lloydia 2:150. ♀.

Biology: Cole, 1953. Tenn. Acad. Sci., Jour. 28:243 (N. Mex.). —Gregg, 1963. Ants of Colo., pp.
298-300.

Myrmica emeryana
***authority mismatch
emeryana Forel. Newfoundland s. to Ga., w. to Man., Idaho, Colo., Ariz. Ecology:

Nests are usually in woodlands in moist, shady situations under stones or other objects.
Sometimes considered as a subspecies of the Palearctic M. schencki Emery.
Myrmica scabrinodis schencki var. emeryana Forel, 1914. Deut. Ent. Ztschr., p. 617. 9,9,

6.
Myrmica schencki latifrons Starcke, 1927. Tijdschr. v. Ent. 70:84. ♀.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:114, 117 (larva). —Smith, 1954. Brooklyn
Ent. Soc, Bui. 49:138-140 9, 9 (worker, female; type - locality). —Wheeler and Wheeler,
1972. Ga. Ent. Soc., Jour. 7:233 (larva).

Biology: Wheeler, 1915. Psyche 22:206. —Talbot, 1945. Ent. Soc. Amer., Ann. 38:365-372
(population studies). — Kannowski, 1959. Insectes Sociaux 6:121. — Medler, 1958. Ent. Soc.
Wash., Proc. 60:258 (swarming). —Peterson and Davies, 1960. Canad. Jour. Zool. 38:15
(predation on Simulium; behavior). — Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
103-104. —Gregg, 1963. Ants of Colo., pp. 311-312, 314.




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Myrmica emeryana
***authority mismatch
tahoensis Wheeler. Mont., Wyo., Utah, Ariz. w. to B. C, Oreg., Nev., Calif. Ecology:
Mostly in mountains at higher elevations.
Myrmica scabrinodis scheyicki var. tahoensis Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:504. 5, 9, d.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7 (3):3^ (Nevada Test Site).

Myrmica hamulata
***authority mismatch
hamulata Weber. Colo., N. Mex., Utah, Ariz. Ecology: Apparently prefers to nest in
upland plateaus from 7,000 to 8,000 ft.
Myrmica sabuleti hamulata Weber, 1939. Lloydia 2:146. 5, 9, 6.

Biology: Gregg, 1963. Ants of Colo., pp. 300-301.

Myrmica hamulata
***authority mismatch
trullicomis Buren. Iowa. The exact status of this form will remain uncertain until a
male can be associated with the workers.
Myrmica sabuleti trullicomis Buren, 1944. Iowa State Col., Jour. Sci. 18:281. 9,9.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool. Bui. 104:99-100.

Myrmica incompleta
***authority mismatch
incompleta Provancher. Labrador s. to N. J., w. to Rocky Mtns., Colo., Utah, N.

Mex. Ecology: Widely distributed with wide elevational tolerance. Prefers moist, grassy

habitats where it usually nests under objects. Host of the inquiline Leptothorax

provancheri Emery.
Myrmica incompleta Provancher, 1881. Nat. Canad. 12:359. 5, 9, 6.
Myrmica rubra brevinodis Emery, 1895. Zool. Jahrb., Abt. f. System 8:312. ♀, ♂.
Myrmica rubra brevinodis var. canadensis Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. 5:76.

9, 9, S.
Myrmica rubra brevinodis var. subalpina Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. 5:77. ♀.

9,6.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:116 (larva). — Francoeur and Beique, 1966.
Canad. Ent. 98:141 (Provancher types).

Biology: Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. 5:73, 77-83. —Wheeler, 1916. Conn. State
Geol. and Nat. Hist. Survey Bui. 22:587. — Kannowski, 1959. Insectes Sociaux 6:121, 155
(pleometrosis). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 99-101. —Gregg, 1963.
Ants of Colo., pp. 291-292, 294.

Myrmica incompleta
***authority mismatch
kuschei Wheeler. Alaska.

Myrmica brevinodis var. kuschei Wheeler, 1917. Harvard Univ., Mus. Comp. Zool., Bui.

61:17. 9, 9.
Myrmica brevinodis var. aUiskensis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.

52:503. ♀.

Myrmica incompleta
***authority mismatch
sulcinodoides Emery. S. Dak., Colo., N. Mex. w. to Alta., Alaska, B. C, Oreg.,

Calif. Some authors do not consider this a valid subspecies.
Myrmica rubra brevinodis var. sulcinodoides Emery, 1895. Zool. Jahrb., Abt. f. System.

8:313. ♀.
Myrmica i^bra brevinodis var. frigida Forel, 1902. Ent. Soc. London, Trans., p. 699. ♀.
Myrmica rubra brevinodis var. whymperi Forel, 1904. Soc. Ent. Belg., Ann. 48:154. ♀.

Biology: Wheeler, 1915. Psyche 22:206. —Cole, 1934. Psyche 41:223. —Gregg, 1963. Ants of
Colo., pp. 293-295, 297.

Myrmica lampra Francoeur
Que. (Pare des Laurentides). Ecology: Apparently parasitic, taken from
nest of another Myrmica. Host: Myrmica sp.
Myrmica lampra Francoeiu", 1968. Nat. Canad. 95:729. ♀, ♂.
lobicomis fracticomis Emery. Newfoundland s. to Tenn., Ohio w. to Rocky Mtns. Colo., N.
Mex., Utah, Ariz. Ecology: Nests are usually near a stream or slough, under stones or
wood. Host of the inquiline Leptothorax provancheri Emery. M. lobicomis lobicomis
Nylander is Palearctic.
Myrmica rubra scabrinodis var. fracticomis Emery, 1895. Zool. Jahrb., Abt. f. System.

8:313. ♀.
Myrmica rubra scabri7iodis var. detrinodis Emery, 1895. Zool. Jahrb., Abt. f. System.
8:316. 9 . A questionable synonym.




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Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:117 (larva). —Cole, 1953. Tenn. Acad. Sci.,
Jour. 28:242-243.

Biology: Wheeler, 1916. Conn. State Geol. and Nat. Hist. Survey Bui. 22:587-588. — Eidmann,
1933. Zool. Anz. 101:203. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:288, 305. —Weber, 1941.
Canad. Ent. 73:140-141 (effect of drouth on nesting habits in prairie states and provinces).
—Weber, 1942. Canad. Ent. 74: 62. — Kannowski, 1957. Psyche 64: 1-5 (host of Leptothorax
provancheri Emery). — Kannowski, 1959. Insectes Sociaux 6:121-124, 155-156 (flight
activities, colony founding). — Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 104-106.
—Gregg, 1963. Ants of Colo., pp. 300, 302-303.
lobicomis lobifrons Pergande. Colo., N. Mex., Utah, Ariz., n. w. to Alaska. Ecology: In

mountains at high elevations. Some authors consider this a synonum of fracticomis.
Myrmica sabuleti var. lobifrons Pergande, 1900. Wash. Acad. Sci., Proc. 2: 521. ♀.
Myrmica rubra scabrinodis var. glacialis Forel, 1904. Soc. Ent. Belg., Ann. 48: 154. ♂.

Biology: Gregg, 1963. Ants of Colo., pp. 303-307.

Myrmica mexicana Wheeler
Ariz., Mexico.

Myrmica mexicana Wheeler, 1914. N. Y. Ent. Soc., Jour. 22: 52. 5, 9, 6.

Taxonomy: Gregg, 1961. N. Y. Ent. Soc, Jour. 69: 211.

Myrmica monticola Wheeler
Que., Mich. w. to Man., N. Dak., Colo. Ecology: Nests in woodlands under
cover of objects.
Myrmica scabrinodis schencki var. monticola Wheeler, 1917. Amer. Acad. Arts and Sci.,

Proc. 52:505. ♀, ♂.
Myrmica sabuleti nearctica Weber, 1939. Lloydia 2: 148. $ , 9 , 6.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:117 (larva).

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 243. —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 106-108. —Gregg, 1963. Ants of Colo., pp. 306-309. — Francoeur,
1966. Nat. Canad. 93: 455 (Que.).

Myrmica pinetorum Wheeler
Mass. s. to S. C, w. to Ohio, Okla., Miss. Ecology: Prefers to nest in sandy
soil.
Myrmica punctiventris pinetorum, Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:384. ♂.
9.

Biology: Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bui. 17:10. —Wesson and Wesson,
1940. Amer. Midland Nat. 24:90, 94.

Myrmica punctiventris Roger
Mass. s. to Ga., w. to Iowa, Nebr., Ark. Ecology: Nests have been found
in soil and rotten logs in woodlands.
Myrmica punctiventris Roger, 1863. Berlin Ent. Ztschr. 7:190. ♂.
Myrmica punctiventris var. isafahani Forel, 1922. Rev. Suisse de Zool. 30:92. ♂, ♀.

Taxonomy: Gregg, 1961. N. Y. Ent. Soc., Jour. 69:211.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:383-384. —Cole, 1940. Amer. Midland
Nat. 24:55. — Headley, 1943. Ohio Jour. Sci. 43:25-26. — Buren, 1944. Iowa State Col., Jour.
Sci. 18:283. —Kannowski, 1959. Insectes Sociaux 6:124.

Myrmica rubra (Linnaeus)
Que., Maine, Mass., R. I.; Europe. Ecology: Unlike other species of Myrmica,

Myrmica rubra
***authority mismatch
is pugnacious and can inflict a painful sting. Probably accidentally introduced

from Europe.
Formica rubra Linnaeus, 1758. Syst. Nat., Ed. 10 1:580.
Myrmica laevinodis Nylander, 1846. Acta Soc. Fenn. 2:927. 5 , 9 , S.
Myrmica rubra champlaini Forel, 1901. Naturhist. Mus. Hamburg Mitt. 18:80. 2 .
Myrmica rubra laevinodis var. bruesi Wheeler, 1906. Psyche 13:38. 5 , 9 , S.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 58:119 (larva). — Picquet, 1958. Dijon Univ.
Lab. de Zool. Trav. 23:28 (larva). —Yarrow, 1955. Roy. Ent. Soc. London, Proc., Ser. B:
Taxonomy. 24:113-115. —Wheeler and Wheeler, 1960. Ent. Soc. Amer., Ann. 53:4-5 (larva).

Biology: Wheeler, 1908. Jour. Econ. Ent. 1:337-339 (introduced into Mass.). — Sturtevant,
1931. Psyche 38:75. —Brian, 1951. Experientia 7:182 (caste determination). —Brian, 1951.




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Physiol. Comp. and Oecol. 2:248-262. —Brian, 1953. Physiol. Comp. and Oecol. 3:25-36
(oviposition by workers). — Brian, 1954. Insectes Sociaux 1:101-122 (caste differentiation,
larval dormancy, winter size, vernalization). — Brian, 1955. Insectes Sociaux 2:1-34 (caste
differentiation, growth of workers and intercastes). — Brian, 1955. Insectes Sociaux
2:85-114 (caste differentiation, larval dormancy, winter size, and vernalization). — Brian
and Brian, 1955. Evolution 9: 280-290 (macrogynes and microgynes). — Brian, 1956.
Insectes Sociaux 3:369-394 (caste differentiation, controlled larval nutrition). — Brian, 1956.
Jour. Anim. Ecol. 25:319-337 (segregation of Myrmica spp.). — Brian, 1957. Insectes
Sociaux 4:191-210 (serial organization of brood). — Brian, 1957. Physiol. Comp. and Oecol.
4:329-345 (food distribution and larval size in cultures). — Weir, 1958. Insectes Sociaux
5:315-339 (polyethism in workers). — Brian, 1962. Insectes Sociaux 9:295-310 (social
conditions affecting early larval differentiation). — Brian, 1963. Insectes Sociaux 10:91-102
(caste differentiation). —Plateaux, 1960. Insectes Sociaux 7: 221. —Brian, 1965. Insectes
Sociaux 12:347 (caste differentiation). — Brian, et al., 1967. Insectes Sociaux 14:13-24
(caste differentiation). —Brian, 1969. Insectes Sociaux 16:249-268 (ecological notes).
Caisson, 1969. Insectes Sociaux 16:279-312. —Brian, 1970. Animal Behavior 18:467-472
(communication between queens and larvae).

Morphology: Tulloch, 1936. Ent. Soc. Amer., Ann. 29:81-84 (metastemal glands). —Allen,
1957. Ent. Monthly Mag. 93:136-139 (intercastes). —Weir, 1959. Insectes Sociaux 6:375-386
(rectro-cerebral endocrine glands and their relation to larval growth and development).

Myrmica spatulata Smith
Tenn., Miss., 111.

Myrmica schencki var. spatulata Smith, 1930. Ent. Soc. Amer., Ann. 23:566. ♂, ♀.

Biology: Smith, 1931. Ent. News 42:21.

Myrmica striolagaster Cole
Tex., Colo., N. Mex., Ariz.

Myrmica stnolagaster Cole, 1953. Tenn. Acad. Sci., Jour. 28:81. ♂.

Taxonomy: Cole, 1957. Tenn. Acad. Sci., Jour. 32:208-209 (male). —Gregg, 1961. N.Y. Ent. Soc,
Jour. 69:211. —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:234 (larva).

Biology: Gregg, 1963. Ants of Colo., pp. 313-314.

Myrmica wheeleri Weber
Ariz. (Mt. Lemmon and Mt. Stratton in Santa Catalina Mtns.).
Myrmica wheeleri Weber, 1939. Lloydia 2:150. 5, 9, 6.

Unplaced Taxon of Myrmica

Myrmica rubra
***authority mismatch
neolaevinodis Forel. "From New York with iris roots". The iris from which the ants

were taken may not have come from New York; consequently, this form may not belong
in the Nearctic fauna.
Myrmica rubra neolaevinodis Forel, 1901. Naturhist. Mus. Hamburg Mitt. 18: 80. ♀.

Genus PARAMYRMICA Cole

Paramyrmica Cole, 1957. Tenn. Acad. Sci., Jour. 32:37^2.
Type-sp>ecies: Paramyrmica colax Cole. Monotypic.

Little is known of the habits of the two species included in this genus. The genus is close to
Myrmica, but differences in the larvae support the distinctness of Paramyrmica.

Taxonomy: Gregg, 1961. N. Y. Ent. Soc., Jour. 69:209-220.

Paramyrmica colax Cole
Tex. (Limpia Canyon, Davis Mtns., Jeff Davis Co.). Ecology: An inquiline. Host:
Myrmica striolagaster Cole.
Paramyrmica colax Cole, 1957. Tenn. Acad. Sci., Jour. 32:37-41. ♂, ♀.

Taxonomy: Wheeler and Wheeler, 1959. Tenn. Acad. Sci., Jour. 34:291-220 (larva).

Paramyrmica rugiventris (Smith)
Colo., Ariz., s. Calif.

Tetramorium rugiventris Smith, 1943. Ent. Soc. Wash., Proc. 45:2, 4. ♂.

Taxonomy: Brown, 1957. Breviora 72:5-7 (an endemic species; in Myrmica). — Gregg, 1961.
N. Y. Ent. Soc., Jour. 69:209 (in Paramyrmica). — Francoeur, 1968. Nat. Canad. 95:728
(probably in Myrmica).




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Geiius MANICA Jurine

Manica Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 276.

Type-species: Formica rubida Latreille. Desig. by Wheeler, 1911.
Aphaenogaster subg. Neomyrma Forel, 1914. Rev. Suisse Zool. 22:275.

Type-species: Aphaenogaster {Neomyrma) calderoni Forel. Monotypic.
Myrmica subg. Oreomyrma Wheeler, 1914. Psyche 21:118.

Type-species: Formica rubida Latreille. Orig. desig.

Four of the five species in this genus are found in western North America, west of the hun-
dredth meridian; the other is Palearctic. Colonies are small and nests are usually found in
openings in coniferous forests commonly under stones in creek or river bottoms. For an excel-
lent account of each species, see Wheeler and Wheeler (1970).

Revision: Wheeler, 1914. Psyche 21:118-122. —Wheeler and Wheeler, 1970. Kans. Ent. Soc,
Jour. 43:129-162.

Taxonomy: Weber, 1947. Ent. Soc. Amer., Ann. 40:439. —Wheeler and Wheeler, 1952. Psyche
59:123 (larvae). —Cole, 1957. Tenn. Acad. Sci., Jour. 32:213 (key to males). —Wheeler and
Wheeler, 1960. Ent. Soc. Wash., Proc. 62:15 (larva). —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 108-110. —Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:234-236 (larvae).

Biology: Wheeler and Wheeler, 1970. Kans. Ent. Soc, Jour. 43:129-162. —Wheeler and
Wheeler, 1970. Kans. Ent. Soc, Jour. 43:363 (additions to natural history of Manica).
—Went, Wheeler, and Wheeler, 1972. BioScience 22:82-88 (feeding and digestion).

Morphology: Fales, et al., 1972. Jour. Ins. Physiol. 18: 1077-1088 (alarm pheromones derived
from the mandibular galnd).

bradleyi (Wheeler). W. Nev., Calif. Ecology: Typically found in openings in coniferous forests
in the Sierra Nevada Mtns. of w. Nev. and Calif., and the Transverse Ranges in s. Calif.
Host of Manica parasitica (Creighton), but the exact relationship is not known.

Myrmica bradleyi Wheeler, 1909. N. Y. Ent. Soc, Jour. 17:77. ♂.

Aphaenogaster (Neomy-niia) calderoni Forel, 1914. Rev. Suisse de Zool. 22:275. ♂.

Taxonomy: Cole, 1957. Tenn. Acad. Sci., Jour. 32:210-212 (female, male). —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62:5-6 (larva). —Wheeler and Wheeler, 1972. Ga. Ent. Soc,
Jour. 7:234-235 (larva).

Biology: Creighton, 1934. Psyche 41:188-189. — Mallis, 1941. South. Calif. Acad. Sci., Bui.
40:65.
hunteri (Wheeler). Alta., Mont., Wyo., Idaho, Utah, B. C, Wash., Oreg., Nev., Calif. Ecology:
Typically found in openings in coniferous forests.
Myrmica (Oreomyrma) hunteri Wheeler, 1914. Psyche 21:119, 121. ♂.
Myrmica (Oreomyrma) aldrichi Wheeler, 1914. Psyche 21:119, 120. ♂.

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31:262. —Cole, 1957. Tenn. Acad. Sci., Jour.
32:212-213 (male). —Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:235 (larva).
mutica (Emery). N. Dak., S. Dak., Colo., N. Mex. w. to Alaska, B. C, Wash., Oreg., Calif.
Ecology: The most widely distributed ant of the genus in N. Amer. It is more
xerophilous and occurs in a greater variety of habitats. Host of the inquiline
Symmyrmica chamberlini Wheeler.
Myrmica mutica Emery, 1895. Zool. Jahrb., Abt. f. System. 8:311. ♂.

Taxonomy: Cole, 1957. Tenn. Acad. Sci., Jour. 32: 213 (male). —Wheeler and Wheeler, 1960.
Ent. Soc Wash., Proc. 62:5-6 (larva). —Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour.
7:235 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:6-7. —Wheeler, 1910. Ants, pp.
432-434. —Cole, 1953. Tenn. Acad. Sci., Jour. 28:243. —Gregg, 1963. Ants of Colo., pp.
314-317. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 109-110.
parasitica (Creighton). Calif. (Alpine Co.; Yosemite Nat. Pk, Mariposa Co.). Ecology:
Collections made from ants nests. Host: Manica bradleyi (AVheeler).
Myrmica (Manica) parasitica Creighton, 1934. Psyche 41:185. ♂.




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Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:66. —Wheeler and Wheeler, 1968.
Pan-Pacific Ent. 44:71-72 (rediscovery).

Genus POGONOMYRMEX Mayr

Members of this genus are collectively known as harvester ants and some species are among
the most conspicuous ants in the arid regions of the western United States and Mexico because
of their mound building habits. The workers collect seeds for food, harvesting plants in their
nesting areas by snipping off seeds with their mandibles. The seeds are stored and are the main
food source though the ants are also scavengers. Nests are in the soil in areas fully exposed to
the sun; some are under stones and others are surmounted by soil craters or by small to huge
mounds with or without coverings of gravel. Some species alter the area surrounding their nest
by clearing away the vegetation. Some species have a painful sting, procure seeds from cul-
tivated crops and damage rangelands.

Revision: Wheeler, 1902. Amer. Nat. 36:85-100. —Wheeler, 1902. Psyche 9:387-393.

—Wheeler, 1914. Psyche 21:151-157. — Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bui.
77:493-514. —Cole. 1968. Pogonomyrmex Harvester Ants, 222 pp.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:106-111 (larvae). —Cole, 1954. Tenn. Acad.
Sci., Jour. 29:117-119. —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:2 (larva).
—Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:236-237 (larvae).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bui. 116:477-478

(proventriculus). — McGurk, et al., 1966. Jour. Ins. Physiol. 12:1435-1441 (identification of
4-methyl-3-heptanone).

Genus POGONOMYRMEX Subgenus POGONOMYRMEX Mayr

Pogonomyrmex Mayr, 1868. Soc. Nat. Modena, Ann. 3:169.

Type-species: Formica badia Latreille. Desig. by Wheeler, 1911.

Species Group Barbatus

Pogonomyrmex anergismus Cole
N. Mex. (15 mi. e. Silver City, 6900 ft.). Ecology: Possibly a social parasite;
found in nest of P. rugosus. Host: Pogonomyrmex ntgosus Emery.
Pogonomyrmex {Pogonomyrmex) anergismus Cole, 1954. Tenn. Acad. Sci., Jour.
29:115-116. ♀, ♂.

Pogonomyrmex apache Wheeler
Southwest. Kans. (?), w. Tex., s. Colo., N. Mex., s. Ariz., s. Nev.; Mexico.
Ecology: Colonies are small and the usually obscure nests are in stony soils with no
superstructure though sometimes in sandy soil with small circular craters.
Pogonomyrmex apache Wheeler, 1902. Psyche 9:392. ♂.
Pogonomyrmex sancti-hyacinthi Wheeler, 1902. Psyche 9:388. ♀.

Biology: Wheeler, 1910. Ants, p. 283. —Cole, 1954. Tenn. Acad. Sci., Jour. 29:266-267.
—Gregg, 1963. Ants of Colo., pp. 318-319.

Pogonomyrmex barbatus (Smith)
E. Ark., e. La., s. and w. Kans., Okla., Tex., e. Colo., s. and central N. Mex.,
s.e. and central Ariz., Nev. (Clark Co.); Mexico. Ecology: The usual nest is a low to high
gravel mound, frequently with a scooped-out center, though the nest structure varies.
The most conspicuous ant of this genus in its range. Red harvester ant.
Myrmica harbata Smith, 1858. Cat. Hym. Brit. Mus. 6:130. ♀.

Myrmica {Atta) molefaciens Buckley, 1860. Acad. Nat. Sci. Phila., Proc. 12:445. ♂, ♀.
Pogonomyrmex barbatiis var. nigrescens Wheeler, 1902. Psyche 9:389, 391. ♂.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:308 (each caste). —Wheeler, 1914. N.
Y. Ent. Soc., Jour. 22:51-52 (each caste). — Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bui. 104:116-119. —Wheeler and Wheeler, 1952. Psyche 59:107 (larva). —Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:2 Oarva).

Biology: McCook, 1879. The Natural History of the Agricultural Ant of Texas, 311 pp.
—Wheeler, 1901. Soc. Ent. de Belg., Ann. 45:202. —Wheeler, 1901. Amer. Nat. 35:723-724.
—Wheeler, 1910. Ants, pp. 11, 85, 179, 197, 202-203, 222, 264, 284, 286-288, 290-293.
—Hunter, 1921. U. S. Dept. Agr. Bur. Ent. Cir. 148:4-7. —Gordon, 1943. Jour. Econ. Ent.




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36:354 (damage to airplane runways). — Michener, 1948. N. Y. Ent. Soc, Jour. 56:239-242.
— Cassidy, et al., 1950. U. S. Dept. Agr. Cir. 842:15. —Barnes and Nemey, 1953. U. S.
Dept. Agr. Farmers' Bui. 1668:1-11. —Young and Howell, 1954. Okla. Acad. Sci., Proc.
35:60-62 (mating swarms). —Young and Gonzalez, 1957. Mex. Secretaria de Agr. y
Ganaderia, Fol. Tec. 23:1-20. —Michener, 1960. Kans. Ent. Soc, Jour. 33:46 (treetop mating
aggregations). —Box, 1960. Ecology 41:381-382. —Gregg, 1963. Ants of Colo., pp. 319-321.
— Johnson, et al., 1969. Biochemical Genetics 3:429-450 (isozyme genotype-environment
relationships in natural populations).

Morphology: Williams and Williams, 1965. Soc. Exp. Biol. Med., Proc. 119:344-346 (venom).
— Benthuysen and Blum, 1974. Ga. Ent. Soc., Jour. 9: 235-238 (quantitative sensitivity of
ant to enantiomers of its alarm pheromone).

bicolor Cole. S. Ariz.; Mexico. Ecology: Nests are in gravelly soil with a low mound of sand
and with a single entrance. Apparently does not clear plants from nest periphery.
Pogonomyrmex (Pogonomyrmex) bicolor Cole, 1968. Pogonomyrmex Harvester Ants, pp.
59-64. 5, 2, 6.

Pogonomyrmex desertorum Wheeler
W. Tex., s. N. Mex., s. Ariz., s.e. CaUf.; Mexico. Ecology: Nests are in
sandy soil in open areas and have circular craters.
Pogonomyrmex desertorum Wheeler, 1902. Psyche 9:387, 390. ♂.

Pogonomyrmex desertorum var. femigineus Olsen, 1934. Harvard Univ., Mus. Comp.
Zool., Bui. 77:496. ♂.

Taxonomy: Cole, 1954. Tenn. Acad. Sci., Jour. 29:119.

Biology: Wheeler, 1910. Ants, p. 283. —Cole, 1934. Ent. Soc. Amer., Ann. 27:399. — Mallis,
1941. South. Calif. Acad. Sci., Bui. 40:64. — Whitford and Ettershank, 1975. Environ. Ent.
4: 689-696 (factors affecting foraging activity).

Pogonomyrmex rugosus Emery
W. Okla., w. Tex., s.e. and s.w. Colo., N. Mex., s. Utah, Ariz., s. and central

Nev., s. Calif.; Mexico. Ecology: Typically, the nest is a flattened gravel disc, though

there is variation in structure.
Pogonomyrmex barbatus rugosus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:309. 5, cJ.
Pogonom,yrmex barbatus var. fuscatus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:309. ♀.
Pogonomyrmex barbattis var. marfensis Wheeler, 1902. Amer. Nat. 36:98. ♂.
Pogonomyrmex similis Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bui. 77:497, 512. ♀.
Pogonomyrmex barbatus curvispinosus Cole, 1936. Ent. News 47:120. ♂.
Pogonomyrmex barbatus spadix Cook, 1953. The Ants of Calif., pp. 98-99, 3 figs. 2 . No

description.

Taxonomy: Wheeler, 1902. Psyche 9:391, 393 (each caste).

Biology: Wheeler, 1902. Psyche 9:393. —Wheeler, 1910. Ants, pp. 284, 290. —Wheeler, 1917.
Psyche 24:178. —Cole, 1937. Ent. News 48:134. —Cole, 1942. Amer. Midland Nat. 28:367.
— Lindquist, 1942. Jour. Econ. Ent. 35:850-853. —Gregg, 1963. Ants of Colo., pp. 322-324,
325, 327. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 50. —Whitford and
Ettershank, 1975. Environ. Ent. 4: 689-696 (factors affecting foraging activity).

Pogonomyrmex tenuispina Forel
S. Calif. (Deep Canyon); Mexico (Baja Calif.).

Pogonomyrmex desertorum, var. tenuispina Forel, 1914. Soc. Vaud. des Sci. Nat., Bui.

50:269. ♂.
Pogonomyrmex dentatus Olsen, 1934. Harvard Univ., Mus. Comp. Zool, Bui. 77:505. ♂.

Taxonomy: Cole, 1968. Pogonomyrmex Harvester Ants, pp. 77-81. —Wheeler and Wheeler,
1973. Ants of Deep Canyon, pp. 50-51 (first U. S. record).

Species Group CKxroENTALis

Pogonomyrmex anzensis Cole
Calif. (Split Mtn., Anza Desert State Park).

Pogonomyrmex (Pogonomyrmex) anzensis Cole, 1968. Pogonomyrmex Harvester Ants,
pp. 87-89. ♀.

Pogonomyrmex brevispinosus Cole
S. Calif., Nev. (Lyon Co.). Ecology: Nests are in compact sandy soil, each
marked by a low circular crater 3 to 8 inches in diameter with a single entrance.
Pogonom^yrmex {Pogonomyrmex) brevispinosus Cole, 1968. Pogonomyrmex Harvester
Ants, pp. 89-94. 5, 9, d.




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Pogonomyrmex occidentalis (Cresson)
Southwest. N. Dak., w. S. Dak., w. Nebr., central and w. Kans., central
and w. Okla., n. Tex., s.e. Mont., Wyo., Colo., N. Mex., s.e. Idaho, Utah, e. and n. Ariz.,
Nev., e. Calif. Ecology: The populous colonies build conical pebble mounds with basal
entrances and peripheral clearings. Workers are pugnacious and are active harvesters,
feeding on and storing large quantities of seeds. Western harvester ant.

Myrmica occidentalis Cresson, 1895. Ent. Soc. Phila., Proc. 4:426. ♂, ♀.

Myrmica seminigra Cresson, 1865. Ent. Soc. Phila., Proc. 4:427. ♂.

Pogonomyrmex opaciceps Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20:971. ♀.

Pogonomyrmex occidentalis ruthveni Gaige, 1914. Biol. Soc. Wash., Proc. 27:93. 5 , 9 , 6.

Pogonomyrmex occidentalis var. utahensis Olsen, 1934. Harvard Univ., Mus. Comp. Zool.,
Bui. 77:498,509. 9, 9,
Taxonomy: Smith, 1953. Brooklyn Ent. See., Bui. 48:131. —Wheeler and Wheeler, 1952.
Psyche 59:110 (larva).

Biology: McCook, 1882. The Honey Ants of the Garden of the Gods, and the Occident Ants
of the American Plains, pp. 123-160. —Dean, 1905. Kans. Acad. Sci., Trans. 19:164-170.
— Headlee and Dean, 1908. Kans. State Col., Bui. 154:165-180. —Wheeler, 1910. Ants, pp.
145, 200, 202-205, 222, 283-284, 290, 291, 426. — Herrick, 1914. Insects Injurious to the
Household and Annoying to Man, pp. 172-173. —Cole, 1934. Ent. News 45:170. —Cole,
1942. Amer. Midland Nat. 28:366. — Knowlton, 1951. Brooklyn Ent. Soc, Bui. 46:75 (loss of
surface area on western rangeland). —Weber, 1959. Ent. News 70:85 (stings; note on
populations). —Chew, 1960. N. Y. Ent. Soc, Jour. 68:81 (colony size and activity).
—Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 118-120. —Gregg, 1963. Ants of Colo.,
pp. 328-334 (as occidentalis comanche). — Spangler, 1968. Kans. Ent. Soc, Jour. 41:318-323
(behavior). — Lavigne, 1969. Ent. Soc. Amer., Ann. 62:1166-1175 (bionomics and nest
structure). —Clark and Camanor, 1973. Biol. Soc Nev., Occ Papers 34:1-6 (use of seed
stores by Heteromyid rodents). — Lavigne and Rogers, 1974. Wyo. Univ. Agr. Exp. Sta.
Sci. Monog. 26: 1-14 (annotated bibliography).

Pogonomyrmex owyheei Cole
Sask., central and w. Mont., n.w. Wyo., Alta., Idaho, n.w. Utah, B. C, Wash.,

Oreg., n. Nev., n.e. Calif. Ecology: Nest structure and habits are similar to those of P.

occidentalis.
PogonoTnyrmex occidentalis owyheei Cole, 1938. Amer. Midland Nat. 19:240. ♂, ♀.

Biology: Cole, 1932. Ohio Jour. Sci. 32:17-20, 133-146, 2Ab-2AG, 533-538 (as occidentalis).
—Cole, 1933. Ent. News 44:16-19 (as occidentalis). —Cole, 1934. Canad. Ent. 66:193-198
(as occidentalis). —Cole, 1934. Ent. News 45:96-101 (as occidentalis). — WiUard and
Crowell, 1965. Jour. Econ. Ent. 58:484-489 (biology; e. Oreg.). —Lavigne and Rogers, 1974.
Wyo. Univ., Agr. Exp. Sta. Sci. Monog. 26: 14-18 (annotated bibliography).

Pogonomyrmex salinus Olsen
Southeast. Oreg., Nev., e. Calif. Ecology: Found mostly in pinyon-juniper areas;
nests are a low bed of gravel with a cleared area surmounted by one or more craterlike
depressions each with one entrance. A docile ant.
Pogonomyrmex salinus Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bui. 77:498, 510. ♂.

Taxonomy: Wheeler and Wheeler, 1972. Ga Ent. Soc, Jour. 7:237 (larva).

Pogonomyrmex subdentatus Mayr
Southwest. Oreg., w. Nev., Calif. Ecology: A timid, inoffensive ant whose
nests are low, irregular beds of sand and gravel with more than one entrance. Mayr's
citation of this form from Conn, is an error.
Pogonomyrmex subdentattis Mayr, 1870. Zool.-Bot. (iesell. Wien, Verb. 20:971. ♂.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:399. — Mallis, 1941. South. Calif. Acad. Sci., Bui.
40:65.

Pogonomyrmex subnitidus Emery
Nev. (?) (Nye Co., Lyon Co.). coastal central and s. Calif.; Mexico. Ecology:
Nests are in sand or loose, sandy soil surmounted by semicircular or circular craters or
marked by irregular beds of sand.
Pogonomyrmex occidentalis var. subnitidus Emery, 1895. Zool. Jahrb., Abt. f. System.
8:310. ♂.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Mallis, 1941. South. Calif. Acad. Sci., Bui.
40:65. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 52.




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Species Group Maricopa

califomicus (Buckley). W. Tex., s. N. Mex., s. Utah., Ariz., Nev., s. Calif.; Mexico. Ecology:
Nests are surmounted by a circular or semicircular crater of loose sand and have a
single entrance. California harvester ant.

Myrmica califomica Buckley, 1867. Ent. Soc. Phila., Proc. 6:336. ♂.

Pogonomyrmex badius var. estebanitcs Pergande, 1893. Calif. Acad. Sci., Proc. 4:33. ♂, ♀.

Pogonomyrmex califomicus longinodis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:311.
5.

Pogonomyrmex califomicus nitratus Cook, 1953. The Ants of Calif., pp. 99-100, 3 figs. 5.

Biology: Wheeler, 1910. Ants, pp. 188-190, 200-201, 284-286, 290-291. — Essig, 1926. Ins. of
West. N. Amer., p. 861. —Cole, 1934. Ent. Soc. Amer., Ann. 27:399. — Michener, 1942. Sci.
Monthly 55:248-258 (history and behavior of a colony). — Michener, 1960. Kans. Ent. Soc,
Jour. 33:46 (treetop mating habit). — McCluskey, 1969. Amer. Zool. 9:566 (flights).
— Erickson, 1972. Ent. Soc. Amer., Ann. 65:57-61 (mark-recapture techniques for
population estimates). — Wheeler and Wheeler, 1973. Ants of Deep Canyon pp. 53-60.
— Whitford and Ettershank, 1975. Environ. Ent. 4: 689-696 (factors affecting foraging
activity).

Morphology: Shapley, 1921. Natl. Acad. Sci., Proc. 6:687-690 (pterergates). — Tullock, 1930.
Psyche 37:61-70.
Comanche Wheeler. Southwest. Ark., w. La., s. Kans., Okla., e. Tex. Ecology: Nests are in

sandy areas in close proximity to post-oak groves and each is marked by a crescentric
or circular crater of sand or sandy soil 3 inches to 2 feet in diameter with a single
entrance.
Pogonom.yrmex occidentalis com,anche Wheeler, 1902. Psyche 9:392. ♂.

Biology: Wheeler, 1910. Ants, pp. 201, 284-285, 292. — Strandtnuinn, 1942. Ent. Soc. Amer.,
Ann. 38:140.

Pogonomyrmex magnacanthus Cole
S. Nev., w. Ariz., s. Calif.; Mexico. Ecology: Nests are in loose, sandy soil
of open deserts with a circular, shallow crater 4 to 6 inches in diameter.
Pogonomyrmex (Pogonomyrmex) magnacanUvus Cole, 1968. Pogonomyrmex Harvester
Ants, pp. 133-137. 5, 9, d.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 60-61.

Pogonomyrmex maricopa Wheeler
W. Tex., s. Colo., N. Mex., s. Utah., s. Nev., Ariz., s.e. Calif.; Mexico.
Ecology: Nests and habits are similar to those of P. califomicus.
Pogonomyrmex califomictLS maricopa Wheeler, 1914. Psyche 21:155. ♂, ♀.
Pogonomyrmex califomicus bamesi Smith, 1929. Ent. Soc. Amer., Ann. 22:546. ♂.
Pogonomyrmex califomicus sinaloanus Olsen, 1934. Harvard Univ., Mus. Comp. Zool.,
Bui. 77:504. ♂.

Taxonomy: Cole, 1954. Tenn. Acad. Sci., Jour. 29:120.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Gregg, 1963. Ants of Colo., pp. 326-328.
— Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 61-62.

Species Group Badius

Pogonomyrmex badius (Latreille)
N. C, S. C, Ga., Fla., Ala., Miss., La. Ecology: Nests are in sand or sandy
soil usually in open woodlands and grassy fields; they construct single or multiple,
flattened, circular sand craters or dome-shaped sand mounds with depressed tops. The
only species of Pogonomyrmex found east of the Mississippi River and the only N.
Amer. polymorphic species. Florida harvester ant.

Formica badia Latreille, 1802. Hist. Nat. Fourmis, p. 238. ♂, ♀.

Myrmica transversa Smith, 1858. Cat. Hym. Brit. Mus. 6:129. ♂.

Atta crudelis Smith, 1858. Cat. Hym. Brit. Mus. 6:170. ♂, ♀.

Myrmica brevipennis Smith, 1858. Cat. Hym. Brit. Mus. 6:130. 6. A questionable syn.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:2 Oarva).




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Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:384-385. —Turner, 1909. Biol. Bui.
17:161-170. —Wheeler, 1910. Ants, pp. 131, 152, 201, 280, 283-285, 292. — Wray, 1938. Ent.
Soc. Amer., Ann. 31:196-200. —Van Pelt, 1953. Tenn. Acad. Sci., Jour. 28:164-168. —Van
Pelt, 1958. Amer. Midland Nat. 59:11-12. —Carter, 1%2. Elisha Mitchell Sci. Soc, Jour.
78:172. — GoUey and Gentry, 1964. Ecology 45:217-225. — Hangartner, Reichson, and
Wilson, 1970. Animal Behavior 18:331-334 (orientation to nest material). — Gentry and
Carlson, 1970. Assoc. Southeast. Biol., Bui. 17:44 (formation and early development).
— Holldobler and Wilson, 1970. Psyche 77:385-399 (recruitment trails). — HoUdobler, 1971.
Science 171:1149-1151 (homing). — Nickle and Neal, 1972. Fla. Ent. 55:65-66 (foraging
behavior). —Morrill, 1972. Fla. Ent. 55:59-60 (tool using behavior). —Morrill, 1975. Ga. Ent.
Soc, Jour. 10: 50-51 (a predator, Apiomerus crassipes crassipes (F.) (Reduviidae)).

Morphology: Wilson, Durlach, and Roth, 1958. Psyche 65:108-114 (chemical releasers of
necrophoric behavior). — Wilson, 1958. Psyche 65:41-51 (chemical releasers of alarm and
digging behavior). — Hermann and Blum, 1967. Ent. Soc. Amer., Ann. 60:661-668
(morphology and histology of poison apparatus). — Blum and Hermann, 1969. Ga. Ent. Soc,
Jour. 4:23-28 (poison gland, functions of the main glandular elements).

Unplaced Taxon of Pogonomyrmex Subgentus Pogonomyrmex

Pogonomyrmex {Pogonomyrmex) califomicus var. Hindleyi Forel, 1914. Soc. Vaud. des Sci.
Nat., Bui. 50:270. 9. Calif. (Escondido).

Genus POGONOMYRMEX Subgenus EPHEBOMYRMEX Wheeler

Pogonom,yrmex subg. Ephebom-yrmex Wheeler, 1902. Psyche 9:390.

Type-species: Pogonomyrmex naegelii Forel. Desig. by Wheeler, 1911.

Biology: Creighton, 1956. Psyche 63:54-56.

Pogonomyrmex huachucanus Wheeler
N. Mex., Ariz. Ecology: Nests are in stony soil, sometimes under
stones, in the open exposed to the sun; there is little or no excavated soil.
Pogonomyrmex hitachucanns Wheeler, 1914. Psyche 21:151. ♂.

Taxonomy: Creighton, 1952. Psyche 59:76-81 (each caste). —Wheeler and Wheeler, 1972. Ga.
Ent. Soc, Jour. 7:237 (larva).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 26:119.
imberbiculus Wheeler. Southwest. Okla., central and w. Tex., s.w. Colo., N. Mex., s. Ariz., s.
Nev., s. Calif.; Mexico. Ecology: Nests are in open or under stones, sometimes with a
small crater.
Pogonomyrmex imberbiculus Wheeler, 1902. Amer. Nat. 36:87, 97. ♂.
Pogonxymyrmex (Ephebomyrmex) townsendi Wheeler, 1909. N. Y. Ent. Soc, Jour. 17:80. ♂.

Taxonomy: Creighton, 1956. Psyche 63:63-64 9. —Wheeler and Wheeler, 1952. Psyche 59:111
(larva). —Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:237 Oarva).

Biology: Wheejer, 1910. Ants, pp. 283-284, 290, 292. -Wheeler, 1917. Psyche 24:178-179.
—Creighton, 1956. Psyche 63:54-56. —Gregg, 1963. Ants of Colo., pp. 334-336. —Wheeler
and Wheeler, 1973. Ants of Deep Canyon, pp. 63-64.

Pogonomyrmex pima Wheeler
S. Ariz.; Mexico. Ecology: Habits are similar to those of P. imberbiculus.
Pogonomyrmex (Ephebomyrmex) pima Wheeler, 1909. N. Y. Ent. Soc, Jour. 17:79. ♂.

Biology: Wheeler, 1910. Ants, p. 283. —Cole, 1934. Ent. Soc. Amer., Ann. 27:399-400.
—Creighton, 1956. Psyche 63:61.

Tribe PHEIDOLINI

Genus STENAMMA Westwood

Stenamma Westwood, 1840. Introduct. Mod. Class. Ins., Sup. 2:83.

Type-species: Stenamma westwoodii Westwood. Monotypic.
Asemorhoptrum Mayr, 1861. Die Europaischen Formiciden, p. 76.

Type-species: Myrmica lippula Nylander. Monotypic.
Theryella Santschi, 1921. Soc d'Hist. Nat. I'Afrique de Nord, Bui. 12:68.




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Type-species: Theryella myops Santschi. Monotypic.

This is a holarctic genus which also extends into the Neotropical Region. Nests are usually in
wooded areas where the small colonies are found in logs, stumps, branches, in soil beneath rocks,
moss, debris, or humus. Workers are apparently predatory, feeding mostly on other arthropods.
Snelling (1973) set up five species groups for the 12 western species of this genus; other North
American species may belong to one of these groups or form new groups.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:297-301. — Forel, 1901. Soc. Ent. de
Belg., Ann. 45:347-348. —Wheeler, 1903. Psyche 10:164-168. —Smith, 1957. Amer. Midland
Nat. 57:133-174 (N. Amer.). —Snelling, 1973. Los Angeles Co. Mas., Contrib. ScL 245:1-38
(western N. Amer.).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:50-52. (larvae).

Biology: Francoeur, 1966. Ent. Soc. Quebec, Ann. 11:115-119 (4 species in Que.).
brevicome (Mayr). N. S., Que. s. to Va., w. to Ont., Minn., Nebr., Colo. (?). Ecology: Prefers to
nest in wooded areas, though sometimes found in meadows, in soil under stones, in
debris, rotting wood, etc.
Aphaenogaster brevicomis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:443, 447-448. ♂.

9.
Stenamma neoarcticum (!) Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:454. 5 (temnle,
male misdet.).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:373, 382. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bui. 22:585. — Kannowski, 1958. Ent. News 69:231-233
(swarming of males). —Gregg, 1963. Ants of Colo., pp. 346-348. —Francoeur, 1966. Ent.
Soc. Quebec, Ann. 11:115-119.
califomicum Snelling. Calif. Ecology: Found in fern humus and oak leaf litter.

Stenamma califomicum, Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:16-18. ♀.
9.

Stenamma carolinense Smith
N. C. (Hoffman, Richmond Co.). Ecology: Collected in sparsely vegetated
sandy soil.
Stenam.mxi carolinense Smith, 1951. Ent. Soc. Wash., Proc. 53:156-158. ♂.

Stenamma chiricahua Snelling
S. Ariz. Ecology: Found in moimtains in shaded creek bed.

Stenamma chiricahiui Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:7-10. ♂, ♀.
S.

Stenamma diecki Emery
Que., Maine s. to N. C, w. to B. C, Wash., Oreg., Calif.; Mexico. Ecology: A
widespread species adapted to various habitats but apparently absent in the southern
states from S. C. to Fla., w. to Tex. Nests are usually in wooded areas in rotting wood
or under objects.
Stenamma westwoodi diecki Emery, 1895. ZooL Jahrb., Abt. f. System. 8:300. 9,9.
Stenamma westwoodi diecki var. im,pressum, Emery, 1895. ZooL Jahrb., Abt. f. System.
8:301. ♀.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:50 Garva). —Snelling, 1973.
Los Angeles Co. Mus., Contrib. Sci. 245:18-21.

Biology: Wheeler and Wheeler, 1944. N. Dak Hist. Quart. 11:244. —Dennis, 1938. Ent. Soc.
Amer., Ann. 31:284, 304. —Cole, 1950. Tenn. Acad. Sci., Jour. 25:297. —Wheeler and
Wheeler, 1963. Ants of N. Dak, pp. 121-122. —Gregg, 1963. Ants of Colo., pp. 350-351, 353.
—Francoeur, 1966. Ent. Soc. Quebec, Ann. 11:115-119. — Lettendre and Pilon, 1972. Nat.
Canad. 99:73-82 (in Que.).

Stenamma dyscheres Snelling
CaUf. Ecology: Found at low to moderate elevations in the Sierra Nevada
foothills. Specimens have been found in pine duff.
Stenamma dyscheres Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:21-25. ♂, ♀.
S.

Stenamma exasperatum Snelling
Calif. (Calaveras Co. and Yosemite Nat. Pk). Ecology: Found under
stones in Sequoia and pine forests.
Stenamma exasperatum Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:28-30. ♀.




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Stenamma foveolocephalum Smith
Miss. (2 mi. s. of Ackerman). Ecology: Specimens collected from
sandy soil on a thinly wooded hillside.
Stenamnm fovolocephala (!) Smith, 1930. Ent. Soc. Amer., Ann. 23:564. ♂.

Biology: Smith, 1931. Ent. News 42:17.

Stenamma heathi Wheeler
Nev., Calif.; Mexico. Ecology: Collected under rocks in pine forests.
Stenamma brevicome heathi Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:410. ♂.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:30-34.

Stenamma huachucanum Smith
Colo., Ariz. Ecology: Collected from under rocks.
Stenamma htiachucanum. Smith, 1957. Amer. Midland Nat. 57:153. ♂.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:34.

Biology: Gregg, 1963. Ants of Colo., pp. 350-351, 353.

Stenamma impar Forel
Que., Mass. s. to N. C, w. to N. Dak., 111., Mo. Ecology: Nests in soil or rotten
wood.
Stenam,ma brevicome impar Forel, 1901. Soc. Ent. de Belg., Ann. 45:347. ♂, ♀.

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., p. 122. — Francoeur, 1966. Ent. Soc.
Quebec, Ann. 11:115-119.

Stenamma meridionale Smith
Va. s. to Ga., w. to 111., Ark. Ecology: Found in wooded areas in soil.

Steiiximma merndioyiale Smith, 1957. Amer. Midland Nat. 57:169-171. ♂, ♀.

Stenamma occidentale Smith
S. Dak., Colo., N. Mex., Idaho, Utah, Ariz., B. C, Wash., Oreg., Calif.;
Mexico. Ecology: Nests in soil under rocks.
Stenamma occidentale Smith, 1957. Amer. Midland Nat. 57:146. 5, 9, S.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:25-28.

Biology: Gregg, 1963. Ants of Colo., pp. 352-354.

Stenamma punctatoventre Snelling
Calif. Ecology: Specimens collected imder a decayed limb in Utter.
Stenamma punctatoventre Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:10-12. ♂.
9,d.

Stenamma schmittii Wheeler
Que., Maine s. to N. C, w. to Minn., Iowa, Mo., Tenn. Ecology: Nests in
wooded areas under stones, rotten wood and litter.
Stenamma brevicome schmittii Wheeler, 1903. Psyche 10:167. ♂.

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:93. —Francoeur, 1966. Ent. Soc.
Quebec, Ann. 11:115-119.

Stenamma sequoiarum Wheeler
Calif, (coastal region from Siskiyou Co. to San Luis Obispo Co.).

Ecology: Specimens have been found under stones in redwood forests and in redwood
litter.
Stenamma brevicome sequoiarum, Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:520.
5, 9.
Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:28.

Stenamma smithi Cole
Idaho, Utah, Nev. Ecology: Collected from sagebrush and juniper duff.
Stenamma smithi Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7 (3):7. ♂.
Stenamma knmvltoni Gregg, 1972. Great Basin Nat. 32:35-39. 9,9.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:12-16.

Stenamma wheelerorum Snelling
Nev. (Mt. Rose, Washoe Co., 8800 ft.).

Stenamma wheelerorum Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:34-35. 2 ,

9.

Genus APHAENOGASTER Mayr

Aphaenogaster Mayr, 1853. Zool.-Bot. Gesell. Wien, Verh. 3:107.

Type-species: Apliaetiogaster sardaiis Mayr. Desig. by Bingham, 1903.
Deromyrma Forel, 1913. Zool. Jahrb., Abt. f. System. 36:772.

Type-species: Aphaenogaster (Ischnomyrmex) swammerdami Forel. Monotypic.
Planimyrma Viehmeyer, 1914. Zool. Jahrb., Abt. f. System. 37:604.

Type-species: Stenamma (Isch7iom.yrmex) loriai Emery. Orig. desig.




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Aphaenogaster subg. Attomyrma Emery, 1915. Accad. delle Sci. dell'Ist. Bologna, Rend. (n.
s.) 19: 70.

Type-species: Formica subterranea Latreille. Orig. desig.
Novomessor Emery, 1915. Accad. delle Sci. dell'Ist. Bologna, Rend. (n. s.) 19:73.

Type-species: Aphaenogaster (IschnomyrTnex) cockerelli Andre. Orig. desig.
Nystalomyrma Wheeler, 1916. Roy. Soc. So. Austral., Trans. 40:215.

Type-species: Myrmica longiceps Smith. Orig. desig.

Most species of this genus nest in soil beneath a covering object but some may be in rotten
logs or arboreal in dead branches. Two species, mariae and tennesseensis are believed to be tem-
porary social parasites but more investigation is needed. The genus is worldwide. The North
American fauna is in need of intensive study.

Revision: Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:443-446. —Emery, 1895. Zool. Jahrb.,
Abt. f. System. 8:301-306. —Wheeler and Wheeler, 1934. Psyche 41:6-12 (treatae and
forms). — Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci., Proc. 69:343-354
(Novomessor). — Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:147-151 (Novomessor).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:54-62 (larvae). — Browm,
1973. In Meggers, et al.. Tropical forest ecosystems in Afr. and S. Amer., pp. 178-185
(generic syn.). — Brown, 1974. Ent. News 85: 45-47 (justification of synonymy of
Novomessor).

Biology: Fellers and Fellers, 1976. Science 192: 70-72 (tool use in 4 species).

Aphaenogaster albisetosa
***authority mismatch
Mayr. Tex., N. Mex., Ariz.; Mexico. Ecology: Foimd in arid plateaus, at elevations of
2,500 to 5,000 feet; makes small crater nests, often under large stones. Omnivorous, but
has a preference for fruits and seeds.
Aphaenogaster albisetosa Mayr, 1886. Zool.-Bot. Gesell. Wien., Verb. 36:443, 446. ♂.
Novomessor cockerelli var. minor Enzmann, 1947. N. Y. Ent. Soc, Jour. 55:148. ♂.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:70 (larva). — Wheeler and
Wheeler, 1960. Ent. Soc. Wash., Proc. 62:8 (larva).

Biology: Wheeler, 1910. Ants, pp. 280-282. —Cole, 1934. Ent. Soc. Amer., Ann. 27:398.
—Creighton, 1955. Psyche 62:89-97.

Aphaenogaster ashmeadi (Emery)
N. C, Tenn. s. to Fla., w. to Mo., Tex.

Stenamma (Aphaenogaster) treatae var. ashmeadi Emery, 1985. Zool. Jahrb., Abt. f.

System. 8:302. ♂.
Aphaenogaster treatae hamedi Wheeler, 1919. Psyche 26:50. ♂.

Taxonomy: Wheeler, 1919. Psyche 26:50. —Smith, 1924. Ent. News 35:50.

Aphaenogaster boulderensis
***authority mismatch
boulderensis Smith. Ariz. (Horseshoe Is., Mead Lake); Nev., Tex. (?). Ecology:
Probably nests in small colonies beneath stones. Possibly confused with A. mutica
Pergande.
Aphaenogaster (Attomyrma) boulderensis Smith, 1941. Great Basin Nat. 2:118, 120. ♂.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7:9.

Aphaenogaster boulderensis
***authority mismatch
smithi Gregg. N. Mex. (Carrizozo).

Aphaenogaster (Attomyrma) boulderensis smithi Gregg, 1949. Ent. Soc. Wash., Proc.
51:171. ♀.

Aphaenogaster cockerelli Andre
Tex., N. Mex., Ariz., Nev., s. Calif.; Mexico. Ecology: Found in arid plateaus
as high as 7,000 ft. Constructs large craters of pebbles under four inches in height with
a single entrance.
Aphaenogaster (Ischnomyrmex) cockerelli Andre, 1893. Rev. de. Ent. 12:150. ♂.
Aphaenogaster sonorae Pergande, 1893. Calif. Acad. Sci., Proc. 4:34. ♂.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:10 (larva). —Wheeler and
Wheeler, 1972. Ga. Ent. Soc., Jour. 7:239 (larva).

Biology: Wheeler, 1910. Ants, pp. 69, 178, 201, 280-282. —Cole, 1934. Ent. Soc. Amer., Ann.
27:397-398. — Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:67. —Cole, 1953. Tenn. Acad.
Sci., Jour. 28:243. —Creighton, 1955. Psyche 62:89-97. — Whitford and Ettershank, 1975.
Environ. Ent. 4: 689-696 (factors affecting foraging activity).




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Morphology: Vick, Drew, et al., 1969. Ent. Soc. Amer., Ann. 62:723-725 (identification of
hydrocarbons).

Aphaenogaster flemingi Smith
N. C. s. to Fla., w. to Ky., La.

Aphaenogaster texmia flemiyigi Smith, 1928. Ent. News 39:275. ♂.
Aphaenogaster texana macrospina Smith, 1934. Ent. Soc. Amer., Ann. 27:386. ♂.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:238 (larva). —Smith, 1957.
Brooklyn Ent. Soc, Bui. 52:113.

Aphaenogaster floridana Smith
N. C. s. to Fla., Ala.

Aphaenogaster (Attomyrma) floridana Smith, 1941. Great Basin Nat. 2:118. ♀.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc 62:7 (larva).

Aphaenogaster fulva Roger
Vt. s. to Fla., w. to Nebr., Colo., La. Ecology: Nests in rotting wood such as logs
and stumps or in soil under stones or other objects. Food consists mostly of live and
dead insects. Temporary host of A. tennesseensis (Mayr) and possibly of A. mariae
Forel.
Aphaenogaster fulva Roger, 1863. Berlin Ent. Ztschr. 7:190. ♂.
Aphaenogaster fulva var. rubida Enzmann, 1947. N. Y. Ent. Soc, Jour. 55:147. ♂.

Biology: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22:586. —Wheeler,
1910. Ants, pp. 81, 83, 206, 448, 453. —Gregg, 1963. Ants of Colo., pp. 336-337, 339. —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326:24-25.

Aphaenogaster huachucana
***authority mismatch
crinimera Cole. N. Mex. (Bandelier Natl. Mon.).

Aphaenogaster {Allomyrma(!)) huachuA:ana crinirnera Cole, 1953. Tenn. Acad. Sci., Jour.
28:82. 5, 9, S.

Aphaenogaster huachucana
***authority mismatch
huachucana Creighton. Colo., Ariz.

Aphaenogaster {Attomyrma) huachucana Creighton, 1934. Psyche 41:189. ♂.

Taxonomy: Creighton, 1951. Psyche 58:89-99 (each caste).

Biology: Gregg, 1963. Ants of Colo., pp. 338-340.

Aphaenogaster lamellidens Mayr
N. Y. s. to Fla., w. to Ul.y Mo., Tex. Ecology: Typically nests in stumps and
logs and feeds on live and dead insects.
Aphaenogaster lamellidejis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:443, 445. 5, 9, i.
Aphaenogaster lamellidens var. nigripes Smith, 1923. Ent. News 34:308. ♂.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24:50, 52.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:383. —Smith, 1918. Ent. News 29:21.
—Smith, 1924. Ent. News 35:51. —Smith, 1928. Ent. News 39:246. —Dennis, 1938. Ent.
Soc. Amer., Ann. 31:285, 304. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:20-21.

Aphaenogaster mariae Forel
N. Y. s. to Fla., w. to Iowa, Kans. Ecology: Probably a temporary parasite. Host:
Aphaenogaster fulva Roger.
Aphaenogaster mariae Forel, 1886. Soc. Ent. de Belg., Bui. 30:41. ♂.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22:585. — Buren,
1944. Iowa State Col., Jour. Sci. 18:284 (female).

Biology: Wheeler, 1910. Ants, pp. 151, 448. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:93.

Aphaenogaster megommata Smith
Oreg., Nev., Ariz., Calif. Ecology: A crepuscular and nocturnal forager.
Aphaenogaster (.Attomyrma) megommatus Smith, 1963. N. Y. Ent. Soc, Jour. 71:244-246.

9.

Taxonomy: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3):9-ll (female, male).
—Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:238 (larva).
miamiana, Wheeler. Fla., Ala., Miss. Ecology: Possibly arboreal, nesting in branches.

Aphaenogaster {Attomyrma) texana var. m.iamiana Wheeler, 1932. N. Y. Ent. Soc, Jour.
40:5. 5, 9, 6.

Aphaenogaster mutica Pergande
Tex. (?); Mexico (Lower Calif.). Ecology: Recorded from Brownsville, Tex.,
but may not occur in U. S.
Aphaenogaster mutica Pergande, 1895. Calif. Acad. Sci., Proc. 5:891. ♂.




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Aphaenogaster patruelis Forel
Calif. (Coastal Islands); Mexico (Lower Calif., Guadeloupe Is., St. Nicholas Is.).
Aphaenogaster patnielis Forel, 1886. Soc. Ent. de Belg., Ann. 30:41. ♀.
Stenamma (Aphaeiiogaster) patruelis bakeri Wheeler, 1904. Amer. Mus. Nat. Hist., Bui.

20:270. ♂.
Aphaenogaster patruelis willowsi Wheeler, 1933. Calif. Acad. Sci., Proc. 21:64. ♂.

Taxonomy: Wheeler, 1934. Pan-Pacific Ent. 10:133.

Aphaenogaster rudis
***authority mismatch
picea (Emery). N. S. s. to N. C. Ecology: Habits are similar to rudis rudis (Emery), but

Aphaenogaster rudis
***authority mismatch
picea is normally found at higher elevations. Temporary host of A. tennesseensis
(Mayr).
Stenamma {Aphaenogaster) fulvum aquia var. piceum. Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:305. 5, 9, J.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:57-60 (larva).

Biology: Wheeler, 1910. Ants, pp. 195, 282, 447-448, 453. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31:286-287, 305. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:94. — Haskins,
1960. N. Y. Ent. Soc, Jour. 68:66-67 (longevity of fertile females).

Aphaenogaster rudis
***authority mismatch
rudis (Emery). Mass. s. to Fla., w. to Wyo., Colo. Ecology: A common and highly

adaptable species, nesting in soil, under stones or logs, in decaying wood, leaf litter,
hollow stems of plants, or under bark at bases of trees. Food consists of insects, seeds,
and pollen of ground nesting bees. Host of temporary ant parasite Aphaenogaster

Aphaenogaster tennesseensis (Mayr)

Stenamma {Aphaenogaster) fulva aquia var. nuie Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:305. 5, 9. .

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:56 (larva).

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:90, 94. —Talbot, 1951. Ent. Soc.
Amer., Ann. 44:302-307. — Brovm, 1958. Psyche 65:39-40 (nest raided by Formica
subintegra Emery). —Gregg, 1963. Ants of Colo., pp. 340-342. — Bobb, 1965, Jour. Econ
Ent. 58:925 (predator of Neodiprion pratti pratti (Dyar)). —Smith, 1965. U. S. Dept., Agr.,
Tech. Bui. 1326:21-22.

Morphology: (Jotwald, 1969. N. Y. (ComeU) Agr. Expt. Sta., Mem. 408:99-112 (mouthparts).
subterranea occidentalis (Emery). B. C. s. to Nev., Calif. A. subterranea subterranea
(Latreille) occurs in the Mediterranean region and central Europe.
Stenam,ma {Aphaenogaster) subterraneum occidentale Emery, 1895. Zool. Jahrb., Abt. f.

System. 8:301. ♀.
Aphaenogaster subterranea borealis Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:412. ♂.
Aphaenogaster subterranea valida var. manni Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:516. ♂.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:363. —Wheeler and Wheeler, 1953. Ent. Soc.
Wash., Proc. 55:61 (larva).

Biology: Mallis, 1941. South. Calif. Acad. Sci., BuL 40:7. — Bruniquel, 1970. Insectes Sociaux
17:245-252 (biology of A. subterranea in Europe).
subterranea valida Wheeler. S. Dak. to B. C, s. to Colo., Utah. Ecology: Prefers to nest under
stones in moist, shady foothill canyons. Intergrades in northern part of range with
subsp. occidentalis.
Aphaenogaster subterranea valida Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:411. ♀.
9,6.

Biology: Gregg, 1947. Colo. Univ., Studies (ser. D) 2:393. —Gregg, 1963. Ants of Colo., pp.
342-344.

Aphaenogaster tennesseensis (Mayr)
Que., Ont. s. to Fla., w. to Minn., S. Dak., Nebr., Kans., Okla. Ecology: In
early stages of colony formation, probably a temporary parasite in ground nests of other
Aphaenogaster species. Lives in ground nests only when females occur in nests of rudis
and picea, otherwise it is exclusively a wood nesting ant. Host: Aphaenogaster fulva
Roger, A. rudis picea (Emery), A. rudis rudis (Emery).

Atta Tenneseensis (!) Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12:743. ♀.

Atta laevis Mayr, 1862. Zool.-Bot. GeselL Wien, Verh. 12:743. ♀.




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Myrmica subruba Buckley, 1867. Ent. Soc. Phila., Proc. 6:336. 5, "9" = cJ.
Stenamma (Aphaenogaster) tenesseense (!) var. ecalcaratum Emery, 1895. Zool. Jahrb.,
Abt. f. System. 8:301. ♂.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22:585-586.
—Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:61-62 (larva). — Francoeur and
Beique, 1968. Nat. Canad. 95:227 (distribution).

Biology: Wheeler, 1910. Ants, pp. 114, 447-448, 450. —Dennis, 1938. Ent. Soc. Amer., Ann.
31:287-288. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:90, 94. —Smith, 1965. U. S.
Dept. Agr., Tech. Bui. 1326:23-24.

Aphaenogaster texana
***authority mismatch
carolinensis Wheeler. S. Va. to n. Ala. Ecology: Nests are in both wood and soil.

Aphaenogaster texana var. carolinensis Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:414.
5, 9-
Taxonomy: Smith, 1931. Ent News 42:17.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:287, 305.

Aphaenogaster texana
***authority mismatch
texana (Emery). N. C, Tenn. s. to Fla., w. to Kans., Ariz.

Stenamma (Aphaenogaster) fulvum aquia var. texanum Emery, 1895. Zool. Jahrb., Abt. f.

System. 8:306. ♂.
Aphaenogaster texana var. furvescens Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:413.

Aphaenogaster (Deromyrma) Silvestrii Menozzi, 1929. Bol. Lab. Zool. (jen. e Agr. Portici
22:282. ♂, ♀.

Taxonomy: Creighton, 1934. Psyche 41:192. —Wheeler and Wheeler, 1953. Ent. Soc. Wash.,
Proc. 55:62 (larva).

Aphaenogaster treatae
***authority mismatch
pluteicomis Wheeler and Wheeler. Ala. w. to Okla., Tex.

Aphaenogaster treatae pluteicomis Wheeler and Wheeler, 1934. Psyche 41:7, 12. 5, 9, cJ.
Aphaenogaster treatae pluteicomis var. oklahomensis Wheeler and Wheeler, 1934. Psyche
41:10, 12. ♂, ♀.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:62 (larva).

Aphaenogaster treatae
***authority mismatch
treatae Forel. Ont. s. to Fla., w. to Mich., Ohio, 111., Ala. Ecology: Nests are in soil
usually in open areas.
Aphaenogaster treatae Forel, 1886. Soc. Ent. de Belg., Bui. 30:40-41. §, 9, S.
Aphaenogaster treatae wheeleri Mann, 1915. Psyche 22:51. ♂, ♀.

Aphaenogaster treatae pluteicomis var. alabamensis Wheeler and Wheeler, 1934. Psyche
41:10. ♂, ♀.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22:585. —Wheeler,
1919. Psyche 26:50.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:303. —Wheeler, 1910. Ants, pp. 151,
200. —Talbot, 1954. Mich. Univ., Contrib. Lab. Vertebr. Biol. 69:1-9. —Talbot, 1966. Kans.
Ent. Soc, Jour. 39:67-77 (flights).

Aphaenogaster uinta Wheeler
Colo., Idaho, Utah, Nev. Ecology: Nests are in fully exposed areas of great
aridity.
Aphaenogaster uinta Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:517. 5, 9, 6.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:363-364.

Biology: Cole, 1934. Psyche 41:223. —Gregg, 1963. Ants of Colo., pp. 344-346, 347.

Unplaced Taxa of Aphaenogaster

According to Creighton (1950, Harvard Univ., Mus. Comp. Zool., Bui.) the following forms
were described from minims only, and it is not possible to detennine what they actually
represent.

Aphaenogaster aquia (Buckley)
N. Y., Va. In the early literature, this and fulva aquia (Buckley) were used
for what is currently called rudis rudis (Emery).
Myrmica {Monomarium (.')) aquia Buckley, 1867. Ent. Soc. Phila., Proc. 6: 341. ♂, ♀.




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Aphaenogaster fulva
***authority mismatch
aquia var. pusilla (Emery). D. C.

Stenanima (Aphaenogaster) fulvum aquia var. pusillum Emery, 1895. Zool. Jahrb., Abt. f.
System. 8: 306. ♂.

Aphaenogaster texana
***authority mismatch
nana Wheeler. Fla. (Gainesville).

Aphaenogaster (Attomyrma) texana nana Wheeler, 1932. N. Y. Ent. Soc, Jour. 40: 6. ♂.

Aphaenogaster texana
***authority mismatch
punctithorax Cole. Tenn. (Gregoiy's Bald, Great Smoky Mtn. Natl. Pk.).

Aphaenogaster texana punctithorax Cole, 1938. Amer. Midland Nat. 19: 239. ♂.

Genus VEROMESSOR Forel

Novomessor subg. Veromessor Forel, 1917. Soc. Vaud. des Sci. Nat., Bui. 51:235.

Type-species: Aphaetiogaster andrei Mayr. Desig. by Emery, 1921.
Veromessor subg. Lohogyiathus Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:152. Nomen
nudum.

These are harvester ants, most common in the arid Southwest. They feed on seeds and other
Arthropods and usually nest in exposed soil where the nests may or may not be marked with a
small crater or piles of chaff from seeds stored in the nest. They are docile ants which seldom
sting.

Revision: Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci., Proc. 69:354-387.

Taxonomy: Smith, 1956. Pan-Pacific Ent. 32:36-37 (key to workers of U. S. species).

—Wheeler and Wheeler, 1956. Psyche 63:142-143 Garvae). —Wheeler and Wheeler, 1972.
Ga. Ent. Soc., Jour. 7:240 (larvae).

Biology: Creighton, 1953. Amer. Mus. Novitates 1612:1-18. — Went, Wheeler, and Wheeler,
1972. BioScience 22:82-88 (feeding and digestion).

Veromessor andrei (Mayr)
Oreg., Nev., Ariz., Calif.; Mexico. Ecology: Found from sea level up to 3500

feet. Nests are in various types of soil and openings are surrounded by a circular disc or
low, obscure crater.
Aphaenogaster andrei Mayr, 1886. Zool.-Bot. (Jesell. Wien, Verb. 36:443, 448. ♂.
, Veromessor andrei flavus Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci., Proc.
69:361,366. ♀.
Veromessor andrei castaneus Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci.,
Proc. 69:361, 365. ♂.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:306-307 (worker, female). —Wheeler
and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 (larva).

Biology: Wheeler, 1910. Ants, p. 280. —Cole, 1934. Ent. Soc. Amer., Ann. 27:398. — Mallis,
1941. South. Calif. Acad. Sci., Bui. 40:7. — Eckert and MalUs, 1937. Calif. Agr. Expt. Sta.
Cir. 342:24 (economics). —Creighton, 1953. Amer. Mus. Novitates 1612:2-4. — McCluskey,
1958. Science 128:536-537 (daily rhythm of male). —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 65-67.

Veromessor chamberlini (Wheeler)
Calif, (coastal islands and mainland).

Messor chamberlini Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:410. ♂.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 Oarva).

Veromessor lariversi Smith
Nev., Calif. Ecology: Nests are in exposed soU or gravel, and may be marked
by one or more small circular craters several inches in diameter.
Veromessor lariversi Smith, 1951. Great Basin Nat. 11:94-96. ♂.

Taxonomy: Cole, 1955. Tenn. Acad. Sci., Jour. 30:51-52 (female).

Biology: Creighton, 1953. Amer. Mus. Novitates 1612:5-6. —Cole, 1963. Ent. Soc. Amer., Ann.
56:680-682. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3): 12.
lobogrnathus (Andrews). Southwest. N. Dak., Colo., Nev. Ecology: Colonies are populous and
nests are under large stones.
Messor lobognathits Andrews, 1916. Psyche 23:82. ♀.

Taxonomy: Wheeler and Wheeler, 1956. Psyche 63:143-145 (larva). —Cole, 1963. Ent. Soc.
Amer., Ann. 56:680-682 (each caste).




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Biology: Gregg, 1955. Psyche 62:45-52 (rediscovery). — Wheeler and Wheeler, 1956. Psyche
63:140-145 (in N. Dak.). —Wheeler and Wheeler, 1959. Ent. Soc. Amer., Ann. 52:176-179
(behavior, nest structure). — Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 123-125.
—Gregg, 1963. Ants of Colo., pp. 354-356. —Wheeler and Wheeler, 1965, Kans. Ent. Soc,
Jour. 38:55-61. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3):12-13.

Veromessor pergandei
***authority mismatch
(Mayr). Ariz., Nev., Calif.; Mexico. Ecology: Nests are in exposed soil, usually with a
single entrance which is surrounded by a crater of excavated soil. Workers are diurnal
foragers and can remain active during periods of intense heat.
Aphaenogaster pergandei Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:444, 448. ♂.

Taxonomy: Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342:8, 25. —Cole, 1963. Ent.
Soc. Amer., Ann. 56:681. —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 (larva).

Biology: Wheeler, 1910. Ants, pp. 16, 280. —Cole, 1937. Ent. News 48:101. —Mallis, 1941.
South. Calif. Acad. Sci., Bui. 40:68. — Creighton, 1953. Amer. Mus. Novitates 1612:7-17.
—Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3): 13-14. —Wheeler and Wheeler,
1965. Kans. Ent. Soc, Jour. 38:55 (habitat, harvesting habits, defense). —Wheeler and
Wheeler, 1967. Kans. Ent. Soc. Jour. 40:238 (distribution and habitat). — Tevis, 1958.
Ecology 39:695-704 (interrelations vrith some desert ephemerals). — Wheeler and Wheeler,
1973. Ants of Deep Canyon, pp. 67-72. —Clark and Comanor, 1973. Amer. Midland Nat. 90:
467-474 (quantitative examination of spring foraging). — Wheeler and Kissing, 1975.
Pan-Pacific Ent. 51: 205-216 (nest). —Wheeler and Kissing, 1975. Pan-Pacific Ent. 51:
303-314 (behavior). —Kissing and Wheeler, 1976. Pan-Pacific Ent. 52: 63-72 (foraging
responses to changes in seed production).

Veromessor smithi Cole
Nev. (Mercury, Nye Co.), Ariz. Ecology: Nests are in fully exposed areas, each
with a small circular crater of soil about their entrance.
Veromessor smithi Cole, 1963. Ent. Soc. Amer., Ann. 56:678-680. 5, 9, 3.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:240 (larva).

Biology: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3): 14.

Veromessor stoddardi
***authority mismatch
chicoensis Smith. Calif. (Chico). Ecology: The type series was found beneath a large
stone.
Veromessor stoddardi chicoensis Smith, 1956. Pan-Pacific Ent. 32:36-38. ♂.

Veromessor stoddardi
***authority mismatch
stoddardi (Emery). S. Calif.; Mexico. Ecology: Nests have been found in hard clay
soil, each with several openings situated at the bottom of a shallow depression and
without excavated soil.
Stenamma {Messor) stoddardi Emery, 1895. Zool. Jahrb., Abt. f. System. 8:307. ♀.

Biology: Wheeler, 1910. Ants, p. 280. —Creighton, 1953. Amer. Mus. Novitates 1612:17-18.

Genus PHEIDOLE Westwood

Pheidole Westwood, 1841. Ann. and Mag. Nat. ffist. 6:87.

Type-species: Atta providens Sykes. Monotypic
Oecophthora Heer, 1852. Naturf. Gesell. in Zurich, Neujahrsbl. 54:11, 15.

Type-species: Oecophthora pusilla Heer. Monotypic.
Ischnomyrmex Mayr, 1862. Zool.-Bot. Gesell. Wien, Verb. 12:738.

Type-species: Myrmica longipes Smith. Monotypic
Leptomyrma Motschulsky, 1863. Soc. Imp. Nat. Moscou, Bui. 36:17.

Type-species: Leptxymyrma gracilipes Motschulsky. Monotypic.
Pheidolacanthinus Smith, 1864. Linn. Soc. London, Jour., Zool. 8:75.

Type-species: PheidolacanthivMs armattis Smith. Monotypic.
Pheidole subg. Ceratopheidole Pergande, 1895. Calif. Acad. Sci., Proc 5:889.

Type-species: Pheidole (Ceratopheidole) granulata Pergande. Monotypic.
Phidole Bingham, 1903. Fauna Brit. India, Hym. 2:220. Variant spelling.
Epipheidole Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19:664.

Type-species: Epipheidole inquilitia Wheeler. Monotypic.
Sympheidole Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:1, 7.

Type-sf)ecies: Sympheidole elecebra Wheeler. Monotypic.
Phidola Schulz, 1906. Spolia Hym., p. 155. Emend.




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Allopheidole Forel, 1912. Soc. Ent. de Belg, Mem. 19:237.

Type-species: Pheidole kingi Andre. Desig. by Wheeler, 1913.
Decapheidole Forel, 1912. Soc. Ent. de Belg., Mem. 19:237.

Type-species: Pheidole perpusilla Emery. Desig. by Wheeler, 1913.
Isopheidole Forel, 1912. Rev. Suisse de Zool. 20:765.

Type-species: Myrmica longipes Smith. Monotypic.
Elasmopheidole Forel, 1913. Zool. Jahrb., Abt. f. System. 36:43.

Type-species: Pheidole aberans Mayr. Desig. by Emery, 1922.
Cardiopheidole Wheeler, 1914. N. Y. Ent. Soc, Jour. 22:48-51.

Type-species: Pheidole vasliti Pergande. Orig. desig.
Anergatides Wasmann, 1915. Ent. Mitt. 4:281. Uncertain syn.

Type-species: Anergatides kohli Wasmann. Monotypic.
Macropheidole Emery, 1915. Soc. Ent. de France, Bui. p. 190.

Type-species: Pheidole fimbriata Roger. Monotypic.
Scrobopheidole Emery, 1915. Soc. Ent. de France, Bui. p. 190.

Type-species: Pheidole scrobifera Emery. Monotypic.
Stegopheidole Emery, 1915. Soc. Ent. de France, Bui. p. 190.

Type-species: Pheidole (Elasmopheidole) upeneci Forel. Monotypic.
Trachypheidole Emery, 1915. Soc. Ent. de France, Bui. p. 190.

Type-species: Pheidole bicomis Forel. Orig. desig.
Parapheidole Emery, 1915. R. Accad. delle Sci. dell'Ist. Bologna, p. 68.

Type-species: Aphaenogaster oculata Emery. Monotypic.
Pheidole subg. E lectropheidole Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bui. 64:438.

Type-species: Pheidole (E lectropheidole) roosevelti Mann. Desig. by Donisthorpe,
1943.
Bruchomyrma Santschi, 1922. Soc. Cient. Argentina, An. 94:248. Uncertain syn.

Type-si>ecies: Bruchomyrma acutidens Santschi. Monotypic.
Cephalomorium Forel, 1922. Rev. Suisse de Zool. 30:91.

Type-species: Tetramorium, (Cephalomorium,) bahai Forel. Monotypic.
Hendecapheidole Wheeler, 1922. Amer. Mus. Novitates 96:3.

Type-species: Pheidole tachigaliae Wheeler. Orig. desig.
Gallardomyrma Bruch, 1932. La Plata Mus., Rev. 33:271.

Type-species: Gallardxymyrma argentina Bruch. Orig. desig.
Conothorax Karawajew, 1935. Treubia 15:75. Preocc. by Jekel, 1854.

Type-species: Conothorax bilobum Karawajew. Monotypic.
Conothomcoides Strand, 1935. Folia Zool. Hydrobiol. 8:176. N. name for Conothorax

Karavaiev.
Eriopheidole Kusnezov, 1952. Mus. de Entre Rios, Mem. 29:5.

Type-species: Eriopheidole symbiotica Kusnezov. Orig. desig.

A large genus of harvesting ants for which most sjjecies are found in the arid Southwest. Most
species have a dimorphic worker caste with majors or soldiers and minors which are not con-
nected by intermediate forms. Only a few species are polymorphic with intermediate sized wor-
kers. The soldier is characteristic in having an enormous head, way out of proptortion to the rest
of the body, and may function as a seed husker.

Most species harvest seeds for food and chaff piles are sometimes found around the entrances
to the nests. Nests may have craterlike excavations when in exposed soil, or nests may be under
stones or other objects and sometimes in rotten wood. Though the main diet normally consists of
seeds, most species also feed on other animal food. Most colonies are small with 300 or fewer in-
dividuals.

This is a huge cosmopolitan genus for which few satisfactory subdivisions have been made. A
few species had been segregated into subgenera or closely related genera, but the bulk of the
species always remained in the subgenus Pheidole. Emery (1921) attempted a group arrange-
ment for this large subgenus, but this has met with little acceptance in recent years. It is now
the trend to thrust all species into a single imit until satisfactory divisions can be made. The ar-
rangement of this genus essentially follows the generic synonomy by Brown (1973) and the
treatment of the genus by Gregg (1958). >•




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Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20:979-989. — Mayr, 1887. Zool.-Bot.
Gesell. Wien, Verb. 37:582-608. —Emery, 1895. Zool. Jabrb., Abt. f. System. 8:288-297.
— Wbeeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:431-478. —Gregg, 1958. N. Y. Ent. Soc,
Jour. 66:7-48 (key to and list of species).

Taxonomy: Emery, 1921. In Wytsman, Gen. Ins., fasc. 174:77-111 (groups). —Wheeler and
Wbeeler, 1953. Ent. Soc. Wasb., Proc. 55:70-80 (larvae). —Smith, 1953. N. Y. Ent. Soc,
Jour. 61:143 (Ceratopheidole). — Creigbton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser.
3:1-46. —Creigbton, 1957. N. Y. Ent. Soc, Jour. 65:203-212 (vasliti complex). —Cole, 1965.
Ent. Soc. Amer., Ann. 58:173-175 {Epipheidole). —Brown, 1973. In Meggers, et al.,
Tropical forest ecosystems in Afr. and S. Amer., pp. 178-185 (generic syn.).

Biology: Weber, 1948. Ent. News 59:31-35 (food of larvae and adults). — Szlep-Fessel, 1970.
Insectes Sociaux 17:233-244 (regulatory mechanism in mass foraging and recruitment of
soldiers).

Pheidole anastasii Emery
S. Fla.; Mexico, Central Amer. Probably introduced. Reported from
greenhouses in various parts of the East, but only established in southern Fla.
Pheidole anastasii Emery, 1896. Soc. Ent. Ital., Bol. 28:76. 3, %.

Taxonomy: Forel, 1901. Naturhist. Mus. Hamburg Mitt. 18:78 (female).

Biology: Wheeler, 1932. N. Y. Ent. Soc, Jour. 40:6. -Smith, 1933. Fla. Ent. 17:23.

Pheidole bahai (Forel)
N. C. (Faisons).

Tetramonum {Cephalomorium) bahai Forel, 1922. Rev. Suisse de Zool. 30:91. ♂.

Taxonomy: Smith, 1955. Brooklyn Ent. Soc, Bui. 50:99 (correct taxonomic placement).

Pheidole barbata Wheeler
W. Ariz., s. Nev., s.e. Calif. Ecology: In small crater nests in desert areas.
Pheidole barbata Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:448. 5 , %.

Biology: MaUis, 1941. South. Calif. Acad. Sci., Bui. 40:68. -Creigbton and Gregg, 1955. Colo.
Univ. Studies, Biol. Ser. 3:1-3. —Wbeeler and Wheeler, 1973. Ants of Deep Canyon, p. 75.

Pheidole bicarinata
***authority mismatch
bicarinata Mayr. Mich, to Tenn., w. to N. Dak., Wyo., Colo. Ecology: Found in
grasslands where nests are usually under objects.

Pheidole bicarinata Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20:982, 989. %.
Pheidole hayesi Smith, 1924. Ent. News 35:251. 5 , V.

Biology: Hayes, 1925. Ent. News 36:42. —Talbot, 1934. Ecology 15:418, 420, 422. — Buren,
1944. Iowa State Col., Jour. Sci. 18:286. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
126-127. —Gregg, 1963. Ants of Colo., pp. 408-409.

Pheidole bicarinata
***authority mismatch
longula Emery. N. Y., N. Dak., w. Tex., Colo., N. Mex. Ecology: Nests may be in
sand or under stones.
Pheidole vinelandica var. longula Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 292. ♂.
%.

Taxonomy: Wbeeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:453-454. —Cole, 1956. Tenn. Acad.
Sci., Jour. 31:112. —Gregg, 1958. N. Y. Ent. Soc, Jour. 65:37.

Biology: Davis and Bequaert, 1922. Brooklyn Ent. Soc, Bui. 17:9. —Wheeler and Wheeler,
1963. Ants of N. Dak., p. 127. —Gregg, 1963. Ants of Colo., pp. 408, 410.

Pheidole bicarinata
***authority mismatch
paiute Gregg. Nev., s. Calif. Ecology: Nests are beneath stones or with craters.
Pheidole bicarinata paiute Gregg, 1958. N. Y. Ent. Soc, Jour. 66:17-18. 5, %.

Biology: Cole, 1966. Brigbam Young Univ., Sci. Bui., Biol. Ser. 7(3): 15. —Wheeler and
Wheeler, 1973. Ants of Deep Canyon, pp. 80-81

Pheidole bicarinata
***authority mismatch
vinelandica Forel. N. Y. to Fla., w. to N. Dak, Utah, Ariz. Ecology: Nests in rotten
logs, exposed soil, or under objects in grassy areas.
Pheidole bicarinata race vinelandica Forel, 1886. Soc. Ent. de Belg., Ann. 30:45. ^,%, 9,

6.
Pheidole vinelandica laemuscula Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 292. ♂.

■u.

Pheidole vinelandica buccalis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:454. 9, ^, 9.
Pheidole vinelandica longula var. castanea Wheeler, 1915. Amer. Mus. Nat. Hist., Bui.
34:405. 5, %. Preocc. by Smith, 1858.




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Pheidole (Allopheidole) vinelandica var. nebrascensis Forel, 1922. Rev. Suisse de Zool.

30:92. $,T^, 9.
Pheidole vinelandica longula var. huachucana Smith, 1951. In Muesebeck, et al., U. S.

Dept. Agr., Agr. Monog. 2:805. N. name for castanea Wheeler.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc, Jour. 66:18, 36.

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:336-337. —Cole, 1937. Ent. News
48:100. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:92. —Cole, 1953. Tenn. Acad.
Sci., Jour. 28:297. — Reid and Nugara, 1961. Jour. Parasitology 47:885-889 (as intermediate
hosts of a turkey tapew^orm). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 127-128.
—Gregg, 1963. Ants of Colo., pp. 410, 412-413. —Smith, 1965. U. S. Dept. Agr., Tech. Bui.
1326:25-26.
califomica califomica Mayr. Calif. (San Francisco region southwards). Ecology: Nests are
found under objects, in oak galls, or in sandy soil.
Pheidole califomica Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20:981, 984, 987. 5 , %.
Pheidole califomica var. incenata Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:407. 5, %.
Pheidole califomica var. satura Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:407. $ , V.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc, Jour. 7:243 (larva).

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:395-397. — Mallis, 1941. South. Calif. Acad. Sci.,
Bui. 40:68-69.
califomica oregonica Emery. Idaho, Wash., Oreg., n. Calif.

Pheidole oregonica Emery, 1895. Zool. Jahrb., Abt. f. System. 8:288, 291. 2, ^.
Pheidole califomica var. shoshoni Cole, 1933. Ent. Soc. Amer., Ann. 26:618. 5, 'U.
Pheidole califomica var. hagermani Cole, 1936. Canad. Ent. 68:35. 5 , "U.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:19.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:69. —Cole, 1942. Amer. Midland Nat.
28:362.
califomica pyramidensis Emery. Nev. (Pyramid Lake).

Pheidole califomica nevadensis Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:408. 5? , %,

9. Preocc. by Forel, 1901.
Pheidole califomica pyramidensis Emery, 1922. In Wytsman, Gen. Ins., fasc. 174:105. N.
name for nevadensis Wheeler.

Pheidole casta Wheeler
Tex. (Canyon of the Rio Grande, Langtry).

Pheidole casta Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:454. 5, If.

Pheidole cerebrosior Wheeler
N. Mex., Ariz., Calif.; Mexico. Ecology: Nests more frequently in

mountan canyons than on open desert and the species seems to prefer the evergreen
oak association.
Pheidole vinelandica cerebrosior Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:405. 5 , 'H.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:3-5.

Pheidole ceres Wheeler
Tex., Colo., N. Mex., Ariz. Ecology: Host of the permanently parasitic ant P.
elecebra. Parasitized colonies have been found to contain only soldiers and workers of
ceres. Ceres nests under stones in rather dry, sunny localities at altitudes of 5,000 to
9,000 ft. and is a seed-storing form. Parasite: Pheidole elecebra (Wheeler).
Pheidole ceres Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:10. 5, If, 9, cJ.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:20, 35.

Biology: Wheeler, 1919. Amer. Phil. Soc., Proc. 58:25. —Wheeler, 1923. Social Life Among the
Insects, p. 219. —Wheeler, 1910. Ants, pp. 279, 497. —Cole, 1953. Tenn. Acad. Sci., Jour.
28:297. —Gregg, 1963. Ants of Colo., pp. 413-415.

Pheidole clementensis Gregg
Calif. (San Clemente Is., Los Angeles Co.; Orange Co.; San Diego Co.).

Pheidole clementensis Gregg, 1969. Ent. News 80: 93-97. 5, %.

Pheidole clydei Gregg
N. Mex., Ariz., Nev., s. Calif. Ecology: Nests are found in and among rocks and
boulders.
Pheidole (Ceratopheidole) clydei Gregg, 1950. N. Y. Ent. See., Jour. 58: 89. ♂.

Taxonomy: Gregg, 1953. Amer. Mus. Novitates 1673: 1 (soldier, distribution).




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Biology: Creighton, 1964. Psyche 71: 169. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, pp. 84-86.

Pheidole cockerelli Wheeler
Okla., Tex., Colo., N. Mex., Ariz.

Pheidole cockerelli Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 464. 5 , %.

Biology: Gregg, 1963. Ants of Colo., pp. 415-417.
crassicomis crassicomis Emery. N. C, Tenn., Ga. w. to Tex.

Pheidole crassicortiis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 296. If.

Pheidole crassicomis var. diversipilosa Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:467.

9, ■y, 9.

Taxonomy: Forel, 1901. Sec. Ent. de Belg., Ann. 45:350 (male).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:302. —Smith, 1918. Ent. News 29:22.
crassicomis tetra Wheeler. Tex. w. to Ariz.

Pheidole crassicortiis porcula var. tetra Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:467.

Pheidole creightoni Gregg
Oreg., Nev., n. Calif.

Pheidole creightoni Gregg, 1955. Psyche 62:19-28. 5, 9, 6.

Taxonomy: Cole, 1957. N. Y. Ent. Soc., Jour. 65:131 (distribution).

Pheidole davisi Wheeler
N. J. s. to N. C, n. Ala.

Pheidole davisi Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:380. 5, %.

Biology: Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bui. 17:8-9. — Bequaert, 1928. N. Y.
(Cornell) Agr. Expt. Sta., Mem. 101:996.

Pheidole dentata Mayr
Va. to Fla., w. to 111., Kans, Tex. Ecology: Nests of small to large colonies are in
exposed soil with a mound of excavated earth above it, under the cover of objects, or in
rotting wood. Occasionally a house-infesting ant.
Pheidole Morrisi var. dentata Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:457. ^,'H, 6.
Pheidole commutata Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:459. 5, T|.
Pheidole dentata war. faisonsica Forel, 1901. Soc. Ent. de Belg., Ann. 45:352. $, V.
Leptothorax tennesseensis Cole, 1938. Amer. Midland Nat. 19:238. ♀.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:460-461. —Wheeler and Wheeler,
1953. Ent. Soc. Wash., Proc. 55:71 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14:71. —Smith, 1924. Ent. News
35:77. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:281, 304. —Cole, 1940. Amer. Midland Nat.
24:29, 44. —Van Pelt, 1950. Ent. News 61:161-163 (parasitism by an Orasema chalcid).
—Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:27-28. —Wilson, 1975. Science 190: 798-800
(enemy specification in the alarm-recruitment system).

Pheidole dentigula Smith
N. C, Tenn. s. to Fla., w. to La. Ecology: Nests in soil or in well-rotted
stumps.
Pheidole dentigula Smith, 1927. Ent. News 38:310. 9, 'U.

Taxonomy: Smith, 1928. Ent. News 39:245-246 (female). —Cole, 1940. Amer. Midland Nat.
24:42, 45. —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:12 (larva).

Biology: Smith, 1944. Fla Ent. 27:14.

Pheidole desertorum Wheeler
W. Okla., w. Tex., N. Mex., Utah, Ariz., Nev. Ecology: Nests have been
found beneath stones and as small crater mounds in sand.
Pheidole desertorum Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:337. 5, ^, 9, d.
Pheidole deserUrrum var. comanche Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:339. ♂.

•2f, 9.
Pheidole desertorum var. mnricopa Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:339. ♂.
—H.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:362.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:397. —Cole, 1937. Ent. News 48:100. —Gregg,
1963. Ants of Colo., pp. 417-419.




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Pheidole elecebra (Wheeler)
Colo. Ecology: A permanent, workerless parasite in colonies of its host.
Host: Pheidole ceres Wheeler.
Si/Ill piteidole elecebra Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:8. ♀, ♂.

Biology: Wheeler, 1910. Ants, p. 497. —Wheeler, 1923. Social Life Among the Insects, p. 219.
—Gregg, 1963. Ants of Colo., pp. 429-431.

Pheidole flavens Roger
Fla.; W. Indies, Central Amer. Possibly an adventive.
Pheidole flavens Roger, 1863. Berlin. Ent. Ztschr. 7:198. $, 'U.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc, Jour. 66:45-46 (Fla. populations regarded as not
quite typical representatives of Pheidole flavens).

Pheidole floridana
***authority mismatch
constipata Wheeler. Tex. (Austin and New Braunfels).

Pheidole constipata Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:468. 9 , ^, 9 , c^ .

Pheidole floridana
***authority mismatch
floridana Emery. N. C. to Fla., w. to Tex.

Pheidole flavens floridana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 293. >i , %, 9.

Pheidole lauta Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:470. 9, 1|, 9, cJ.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc, Jour. 66:35.

Pheidole gilvescens Wheeler
Ariz., s. Nev., Calif. Ecology: Nests in soil in deserts, usually forms small
craters.
Pheidole xerophila tucsonica var. gilvescens Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.
24:448. $ , "y.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:5-7, 40.

Biology: Cole. 1934. Ent. Soc. Amer., Ann. 27:397. —Cole, 1956. Tenn. Acad. Sci., Jour. 31:113.
— Wlieeler and Wheeler, 1973. Ants of Deep Canyon, p. 76.

Pheidole grundmanni Smith
Utah (Ashley Cr. near Vernal).

Pheidole (Ceratopheidole) grutidmanni Smith, 1953. N. Y. Ent. Soc, Jour. 61:144-146. ♂.

Biology: Gregg, 1963. Ants of Colo., pp. 426-427.

Pheidole humeralis Wheeler
Tex. (Corsicana).

Pheidole humeralis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:456. 5, %.

Pheidole hyatti
***authority mismatch
hyatti Emery. Okla., Tex., Colo. w. to Nev., s. Calif.; Mexico.

Pheidole hyatti Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289-290, 295. 9 , %.
Pheidole hyatti var. ecitonodora Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:463. 9, V,
9, 6.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc 55:74 (larva). — Wheeler and
Wheeler, 1972. Ga. Ent. Soc, Jour. 7:244 (larva).

Biology: Essig, 1926. Ins. of West. N. Amer., p. 859. — Eckert and Mallis, 1937. Calif. Agr.
Expt. Sta. Cir. 342:26. —Gregg, 1963. Ants of Colo., pp. 419-420. —Wheeler and Wheeler,
1973. Ants of Deep Canyon, pp. 79-80.

Pheidole hyatti
***authority mismatch
solitanea Wheeler. Calif, (coastal area in San Diego region).

Pheidole hyatti solitatiea Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:409. ^,%, 9.

Pheidole inquilina (Wheeler)
Nebr., Colo., Nev. Ecology: An inquiline in nests of other ants. Host:
Pheidole pilifera pilifera (Roger), P. pilifera coloradensis Emery.
Epipheidole inquilina Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19:664. Gynandromorph.

Taxonomy: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:15-17 (female, male). —Smith, 1940.
Ent. Soc. Wash., Proc. 42:106-109 (worker). —Cole, 1965. Ent. Soc. Amer., Ann. 58:173-175
(worker, soldier; placement in Pheidole; biological notes).

Biology: Wheeler, 1910. Ants, pp. 107, 113, 150, 156, 497-198. —Wheeler, 1923. Social Life
Among the Insects, pp. 215-219. —Gregg, 1963. Ants of Colo., pp. 427-429. —Cole, 1966.
Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3): 15.

Pheidole lamia Wheeler
Ga., Miss., Tex. Ecology: Apparently hypogaeic, living under stones in small
colonies. The phragmotic head of the soldier is a striking feature of this species.
Pheidole lamia Wheeler, 1901. Amer. Nat. 35:534. 9, %.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:477-478 (soldier, female.).




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Biology: Wheeler, 1910. Ants, pp. 212, 248. —Smith, 1931. Ent. News 42:21-22. —Gregg, 1956.
Ent. News 67:37-39 (also distribution).

Pheidole macclendoni Wheeler
S. Tex. w. to Ariz.

Pheidole macclendoni Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:450. 5 , %.

Taxonomy: Cole, 1957. N. Y. Ent. Soc., Jour. 65:130-131 (worker, soldier). —Gregg, 1958. N.
Y. Ent. Soc., Jour. 66:46-47.

Pheidole marcidula Wheeler
Tex. (Barton Cr., Austin).

Pheidole marcidula Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:457. 9, %.

Pheidole megacephala (Fabricius)
Fla.; W. Indies, Hawaii, and probably all tropical regions of the

world. Introduced, probably native to Africa but spread by commerce throughout the
world. Bigheaded ant.
Formica megacephala Fabricius, 1793. Ent. System. 2:361. %.
Phidola megalocephala Schulz, 1906. Spolia Hym., p. 155. Emend.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:843-844. —Wheeler and Wheeler,
1953. Ent. Soc. Wash., Proc. 55:75 (larva). —Wilson and Taylor, 1967. Pacific Insects
Monog. 14:46-48.

Biology: Illingsworth, 1916. Hawaii. Ent. Soc., Proc. 3:349-368. —Phillips, 1934. (Hawaii
Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bui. 15:5-12. — Broekhuysen, 1948. Union
So. Africa Dept. Agr. Bui. 266:1-40. —Brown, 1958. Acta Hym. 1:47. —Brown, 1959. Bui.
Ent. Res. 50:523. —Weber, 1960. Ent. Soc. Wash., Proc. 62:232. —Taylor and Wilson, 1961.
Psyche 68:143. — Kempf, 1962. Studia Ent. 5:18-19. — Haskins and Haskins, 1965. Ecology
46:737 (competition between P. megacephala and Iridcmiyrmex humilis (Mayr) in
Bermuda). — Fluker, Huddleston, and Beardsley, 1968. Jour. Econ. Ent. 61:474. — Fluker
and Beardsley, 1970. Ent. Soc. Amer., Ann. (53:1290-1296 (sympatric association with
Iridomyrmex humilis and Anoplolepis longipes in Hawaii).
metailescens metallescens Emery. Fla. w. to Tex.

Pheidole metallescens Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 294. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:47^477. —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62:12 Oarva).

Biology: Smith, 1924. Ent. News 34:78 (as splendidula).

Pheidole metallescens
***authority mismatch
splendidula Wheeler. Southwest. Tex.

Pheidole metallescens splendidula Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:474. ♂.
^, 9, 6.

Pheidole micula Wheeler
Ariz., s. Calif. Ecology: Nest was found under a stone.

Pheidole califomica micula Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:408. 5, 'U.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:33. —Wheeler and Wheeler, 1972. Ga. Ent.
Soc., Jour. 7:244 (larva).

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 80.

Pheidole militicida Wheeler
N. Mex., Ariz.

Pheidole militicida Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:398. 5, V.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:9-12 (female, male;
biological notes).

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:395. —Cole, 1953. Tenn. Acad. Sci., Jour. 28:298.
— Creighton and Creighton, 1959. Psyche (56:1-12 (observations on habits in s. Ariz.).

Pheidole moerens Wheeler
Fla., Ala (Mobile); Puerto Rico. Apparently introduced.

Pheidole moerens Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:136-138. 5, V.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:244 Oarva). — Wojcik, Banks,
and Buren, 1975. Coop. Econ. Ins. Rpt. 25 (49-52): 906 (first report in Fla.).

Biology: Smith, 1937. Puerto Rico Univ., Jour. Agr. 20:842-843.

Pheidole morrisi
***authority mismatch
impexa Wheeler. Okla., Tex.

Pheidole morrisi var. impexa Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:461. ^ ,'U, 9 ,
cJ.




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Biology: Lindquist, 1942. Jour. Econ. Ent. 35:850-852.

Pheidole morrisi
***authority mismatch
morrisi Forel. N. Y. to Fla., w. to 111., La., Tex.

Pheidole Morrisii Forel, 1886. Soc. Ent. de Belg., Bui. 30:46. 5 , 1^.

Pheidole morrisi var. Vanceae Forel, 1901. Soc. Ent. de Belg., Ann. 45:351. 9, %, 9, 6.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:461. —Smith, 1924. Ent. News
35:53. —Wheeler and Wheeler, 1960. Ent. Soc. Amer., Ann. 53:12 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:302. —Wheeler, 1905. Amer. Mus.

Nat. Hist., Bui. 21:380. —Davis and Bequaert, 1922. Brooklyn Ent. Soc, Bui. 17:8-9.

Pheidole nuculiceps Wheeler
Tex. (Canal River, New Braunfels).

Pheidole nuculiceps Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:473. 5 , 'H.

Pheidole pilifera
***authority mismatch
artemisia Cole. Utah, Ariz.

Pheidole pilifera artemisia Cole, 1933. Ent. Soc. Amer., Ann. 26:616. 5 , %.

Taxonomy: Cole, 1938. Amer. Midland Nat. 20:372 (female). —Cole, 1952. Tenn. Acad. Sci.,
Jour. 27:280. —Gregg, 1955. Psyche 62:22-23.

Biology: Cole, 1942. Amer. Midland Nat. 28:362.

Pheidole pilifera
***authority mismatch
coloradensis Emery. N. Dak., Colo., N. Mex., Nev. Ecology: A harvester of seeds of
various grasses; nests are usually under stones. Parasite: Pheidole iriquilirM (Wheeler).
Pheidole pilifera var. coloradensis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:288, 291. ♂.

■u.

Pheidole pilifera coloradensis var. neomexicatia Wheeler, 1908. Amer. Mus. Nat. Hist.,
Bui. 24:436. %.

Taxonomy: Cole, 1952. Tenn. Acad. Sci., Jour. 27:279. —Gregg, 1955. Psyche 62:22-23.

Biology: Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19:666. —Wheeler, 1904. Amer. Mus.
Nat. Hist., Bui. 20:16-17. —Wheeler, 1910. Ants, pp. 279, 498. —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 128-130. —Gregg, 1963. Ants of Colo., pp. 419, 421-422. —Cole,
1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7(3): 15.

Pheidole pilifera
***authority mismatch
pacifica Wheeler. Nev., Calif.

Pheidole xerophila pacifica Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:404. 5, "U, 9, S.

Taxonomy: Cole, 1952. Tenn. Acad. Sci., Jour. 27:279. —Gregg, 1955. Psyche 62:22-23.

Pheidole pilifera
***authority mismatch
pilifera (Roger). N. Y., Mass. s. to Fla., w. to N. Dak., Nebr., Kans. Ecology: A

harvester of seeds; most nests have been found in exposed soil and have craterlike
excavations. Parasite: Pheidole inquilina (Wheeler).
Leptotharax pilifer Roger, 1863. Berlin. Ent. Ztschr. 7:180. ♂.
Pheidole pennsylvanica Roger, 1863. Berlin. Ent. Ztschr. 7:199. ♂.
Pheidole pilifera septentrionaiis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:436. %.
Pheidole pilifera var. simulans Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:436. %.

Taxonomy: Cole, 1952. Tenn. Acad. Sci., Jour. 27:280. —Wheeler and Wheeler, 1953. Ent. Soc.
Wash., Proc. 55:79 (larva). —Gregg, 1955. Psyche 62:22-23.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:379-380. —Wheeler, 1910. Ants, pp.
152, 278. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:92. —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 130-131.

Pheidole pinealis Wheeler
Tex.; Mexico. Ecology: Nests have been found under stones.
Pheidole pinealis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:459. 5 , 'U.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:12-15 (also biological
notes).

Pheidole porcula Wheeler
W. Tex., Colo. (?).

Pheidole crassicomis porcula Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:466. $ , "U.

Pheidole psammophila Creighton and Gregg
Ariz., s. Calif.; Mexico. Ecology: Found in sandy areas at
low elevations.
Pheidole psammophila Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:15-19.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 81.




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Pheidole rhea Wheeler
S. Ariz.; Mexico. Ecology: Prefers to nest on plateaus or in foothills at the base
of mountains, 3700 to 7000 ft.
Pheidole rhea Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:452. ♀.

Taxonomy: Smith, 1943. Ent. Soc. Wash., Proc. 45:5-9 (worker, soldier). —Gregg, 1949.
Psyche 56:70-73.

Biology: Gregg, 1949. Psyche 56:70-73.

Pheidole ridicula Wheeler
Tex. (Brownsville area).

Pheidole ridicula Wheeler, 1916. New England Zooi. Club, Proc. 6:29. %.

Biology: Creighton, 1966. Psyche 73:1.

Pheidole rugulosa Gregg
Tex., Ariz.

Pheidole rugulosa Gregg, 1958. N. Y. Ent. Soc., Jour. 66:26-29. ^,'n,9,S.

Pheidole sciara Cole
Tex., N. Mex.

Pheidole sciara Cole, 1955. Tenn. Acad. Sci., Jour. 30:47-49. 5 , %.

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31:116.

Pheidole sciophila Wheeler
Tex. to s. Ariz.; Mexico.

Pheidole sci(yphila Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:443. ^,%,9, 6.
Pheidole jrroserpina Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:437. 9, V.
Pheidole sciophila var. semilaevicephala Smith, 1934. Ent. Soc. Amer., Ann. 27:385. %.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:19-22.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:397.

Pheidole senex Gregg
Tex., Colo., N. Mex.

Pheidole senex Gregg, 1952. Amer. Mus. Novitates 1557:1-4. $, 'U.
Pheidole pilifera anfracta Cole, 1952. Tenn. Acad. Sci., Jour. 27:278. 5 , %.

Taxonomy: Gregg, 1955. Psyche 62:22-23.

Biology: Gregg, 1963. Ants of Colo., pp. 422-424.

Pheidole sitarches
***authority mismatch
campestris Wheeler. Miss., Mo. w. to Colo., central Tex.

Pheidole sitarches rufescens var. campestris Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.

24:443. ?, 'U.
Pheidole sitarches rufescens Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:443. 9, "U, 9.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:37.

Biology: Smith, 1924. Ent. News 35:53. —Gregg, 1963. Ants of Colo., pp. 424-425.

Pheidole sitarches
***authority mismatch
littoralis Cole. Fla. (Lido Beach, Sarasota).

Pheidole sitarches littoralis Cole, 1952. Ent. Soc. Amer., Ann. 45:443-444. $ , "U.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:39.

Pheidole sitarches
***authority mismatch
sitarches Wheeler. Tex. (Brownsville area); Mexico.

Pheidole sitarches Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:440. 5, "y, 9.

Pheidole sitarches var. transvarians Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:442. ♀.

v.

Taxonomy: Cole, 1952. Ent. Soc. Amer., Ann. 45:444. —Gregg, 1958. N. Y. Ent. Soc., Jour.

66:39.
Biology: Wheeler, 1910. Ants, p. 279. —Wilson, 1957. Psyche 64:46-50 (nuptial flights).

Pheidole sitarches
***authority mismatch
soritis Wheeler. W. Tex., N. Mex., Utah, Ariz.; Mexico.

Pheidole soritis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:439. 5, 'H.
Pheidole tepicana cavigenis Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:403. %.

Taxonomy: Cole, 1952. Ent. Soc. Amer., Ann. 45:444. —Gregg, 1958. N. Y. Ent. Soc., Jour.
66:37.

Pheidole spadonia Wheeler
S. Ariz.; Mexico.

Pheidole spadonia Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:400. 9 , 'U.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:22-24 (also biological
notes).




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Pheidole subdentata Pergande
Ariz.; Mexico.

Pheidole subdentata Pergande, 1895. Calif. Acad. Sci., Proc. 5:888. ♀.
Pheidole obtusospinosa Pergande, 1895. Calif. Acad. Sci., Proc. 5:889. 'H.
Pheidole arizonica Santschi, 1909. Soc. Ent. Ital., Bol. 41:3. %.

Taxonomy: Creighton, 1957. N. Y. Ent. Soc, Jour. 65:211.

Pheidole tepicana Pergande
Tex., Ariz.; Mexico. A polymorphic species with the largest soldiers and

smallest workers connected by intermediates, unlike the dimorphic worker castes of

most Pheidole.
Pheidole tepicana Pergande, 1895. Calif. Acad. Sci., Proc. 5:878. $ , 'U.
Pheidole rugifrons Pergande, 1895. Calif. Acad. Sci., Proc. 5:880. %.
Pheidole carbonaria Pergande, 1895. Calif. Acad. Sci., Proc. 5:881. 5 , %.
Pheidole Kingi Andre, 1898. Soc. Ent. de France, Bui. p. 244. § , %.
Pheidole Tawnsendi Andre, 1898. Soc. Ent. de France, BuL, p. 246. 5 , 'U.
Pheidole kingi instabilis Emery, 1901. Soc. Ent. de France, Bui., p. 129. 5 , 'U.
Pheidole kingi torpescens Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:404. 5, 11.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:74 (larva). —Creighton and
Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:24-35.

Biology: Wheeler, 1901. Soc. Ent. de Belg., Ann. 45:203. —Wheeler, 1907. Amer. Mus. Nat.
Hist., Bui. 23:2-12. —Cole, 1956. Tenn. Acad. Sci., Jour. 31:113.

Morphology: Wheeler, 1910. Ants, p. 56.

Pheidole texana Wheeler
Tex. (Travis Co.).

Pheidole texana Wheeler, 1903. Psyche 10:95. 5 , H.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:464.

Pheidole titanis Wheeler
W. Tex. to s. Ariz. Ecology: Diet may be restricted to termites.
Pheidole titanis Wheeler, 1903. Psyche 10:95. 5 , V.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:461-462. —Creighton and Gregg,
1955. Colo. Univ. Studies, Biol. Ser. 3:35-37 (also biological notes; termite feeding).

Pheidole tysoni Forel
Va., N. C, Ohio, Tenn.

Pheidole tysoni Forel, 1901. Soc. Ent. de Belg., Ann. 45:348. 5, "U, d.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24:42, 45.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:282, 304. —Wesson and Wesson, 1940. Amer.

Midland Nat. 24:92.

Pheidole vallicola Wheeler
S. Ariz.

Pheidole cmssicomis vallicola Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:409. 9 , V.

Pheidole virago Wheeler
Tex., Ariz.

Pheidole virago Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:401. 5, ^.

Pheidole vistana Forel
S. Calif.; Mexico. Ecology: Found in deserts in small crater nests in the soU.

Pheidole stisannne longipes Pergande, 1895. CaHf. Acad. Sci., Proc. 5:885. 5 , "U. Preocc. by
Latreille, 1802; Smith, 1858.

Pheidole longipes var. vistana Forel, 1914. Soc. Vaud. des Sci. Nat., Bui. 50:272. ♀.

Pheidole grallipes Wheeler, 1916. Psyche 23:40. N. name for longipes Pergande.

Taxonomy: Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:397-398.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:69. —Wheeler and Wheeler, 1973. Ants
of Deep Canyon, pp. 77-79.

Pheidole xerophila
***authority mismatch
tucsonica Wheeler. Tex., N. Mex., Ariz., Calif.; Mexico. Ecology: Nests in soU may
be craterlike or with small mounds.
Pheidole xerophila tucsonica Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:448. $ , H.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:40-42.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:397. —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 82-83.




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Pheidole xerophila
***authority mismatch
xerophila Wheeler. Tex., N. Mex., s. Calif. Ecology: Found in small crater nests in
the desert soil. The range of this subspecies lies slightly to the south of that of

Pheidole tucsonica
***authority mismatch
in Tex. and N. Mex.
Pheidole xerophila Wheeler, 1908. Amer. Mas. Nat. Hist., Bui. 24:446. ^,'U, i.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:40-42.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 82.

Pheidole yaqui Creighton and Gregg
S. Calif.; Mexico. Ecology: Found in deserts in small crater nests
in the soil.
Pheidole yaqui Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:43-46. $ , %.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 83-84.

Tribe CARDIOCONDYLINI
Genus CARDIOCONDYLA Emery

Cardiocondyla Emery, 1869. Accad. degli Aspiranti Naples, Ann. 2:20.

Type-species: Cardiocondyla elegans Emery. Monotypic.
Emeryia Forel, 1890. Soc. Ent. de Belg. Ann. (C. R.) 34:110.

Type-species: Emeryia wroughtoni Forel. Monotypic.
Xenometra Emery, 1917. Soc. Ent. de France, Bui., p. 96.

Type-sjjecies: Xenometra monilicomis Emery. Monotypic.
Cardiocondyla subg. Dyclona Santschi, 1930. Rev. Suisse de Zool. 37:70. Syn. questionable.

Type-sp)ecies: Monomorium cristatum Santschi. Orig. desig.
Cardiocondyla subg. Loncyda Santschi, 1930. Rev. Suisse de Zool. 37:70. Syn. questionable.

Type-species: Cardiocondyla {Loncyda) monardi Santschi. Monotypic.
Cardiocondyla subg. Prosopidris Wheeler, 1935. Psyche 42:40.

Type-sp)ecies: Cardiocondyla {Prosopidris) sima Wheeler. Orig. desig.

The majority of species of this genus are found in the warmer parts of Europe, Asia, and
Africa. In the United States, members of this genus are found in the southern tier of states
from Florida to California. The colonies are small and nests are constructed in soil and in plant
cavities. They are easily transported by commerce and aU the species below may have been in-
troduced.

Revision: Smith, 1944. Ent. Soc. Wash., Proc. 46:3041 (U. S. species).

Taxonomy: Weber, 1952. Amer. Mus. Novitates 1548:5. —Brown, 1973. In Meggers, et al..
Tropical forest ecosystems in Afr. and S. Amer., pp. 161-185 (generic syn.).

Biology: Wilson, 1960. Psyche 66:29-34 (tandem running).

Cardiocondyla ectopia Snelling
Ariz., s. Calif. Possibly of Old World origin.

Cardiocondyla ectopia Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 76-81, figs. 5, 9, cJ.

Biology: Creighton and Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 87-91 (behavior).

Cardiocondyla emeryi
***authority mismatch
ForeL Fla., Tex.; W. Indies, Africa, Asia, Pacific Islands. Probably introduced. A tramp
species distributed by commerce throughout the tropics.
Cardiocondyla emeryi Forel, 1881. Munchen Ent. Ver., Mitt. 5:5. ♂.

Cardiocondyla nuda subsp. nereis Wheeler, 1927. Amer. Acad. Arts and Sci., Proc. 62:140.
5. 9-
Taxonomy: Borgmeier, 1937. Rev. de Ent. 7:133 (ergatoid male), —Weber, 1952. Amer. Mus.
Novitates 1548:5. —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:53 (Polynesia).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:89. — Snuth, 1936. Puerto Rico Univ.,
Jour. Agr. 20:835. —Creighton and Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 82-86
(behavior).

Cardiocondyla nuda (Mayr)
Fla., Ga., La., Tex.; Pacific Islands. Probably introduced into U. S. from the
Oriental Region.
Leptotharax nudus Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. Kl., Sitzber. 53:508. ♂.
Cardiocondyla nuda war. minutior Forel, 1899. Fauna Hawaiiensis, 1:120. ♀.




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Taxonomy: Wilson and Taylor, 1967. Pacific Ins. Monog. 15:55 (Polynesia). — Wheeler and
Wheeler, 1973. Ga. Ent. Soc, Jour. 8:27 (larva).

Biology: Phillips, 1934. (Hawaii Univ.) Expt. Sta., Pineapple Prod. Coop. Assoc. Bui. 15:22.
— Creighton and Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 86-87 (behavior).

Cardiocondyla venustula Wheeler
Fla., La.; Puerto Rico, Haiti. Probably introduced.

Cardiocondyla venustula Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:128. ♂, ♀.

Biology: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:836. —Smith, 1944. Fla. Ent. 27:15.

Cardiocondyla wroughtonii (Forel)
Fla., Ga.; Oriental Region. Probably introduced.

Emeryia wroughtonii Forel, 1890. Soc. Ent. de Belg., Ann. (C. R.) 34:110. "5" =

ergatomorphic cJ.
Cardiocondyla wroughtonii var. hawaiensis (!) Forel, 1899. Fauna Hawaiiensis 1:119. ♂.
Cardiocondyla urroughtoni var. bimaculata Wheeler, 1929. Lab. Zool. Gen. e Agr. Portici,
Bol. 24:43. $.9.

Taxonomy: Wilson and Taylor, 1967. Pacific Ins. Monog. 14:56 (Polynesia).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:7. —Smith, 1933. Fla. Ent. 17:24.

Tribe CREMATOGASTRINI

Genus CREMATOGASTER Lund

Crematogaster Lund, 1831. Ann. des Sci. Nat., Zool. 23:132.

Type-species: Formica scutellaris Olivier. Desig. by Bingham, 1903.
Cremastogaster Agassiz, 1846. Nomencl. Zool., Index Univ., p. 103. Emend.
Acrocoelia Mayr, 1852. Zool.-Bot. Gesell. Wien, Verb. 2:147.

Type-species: Acrocoelia ruficeps Mayr. Desig. by Wheeler, 1911.
Crematogaster subg. Oxygyne Forel, 1901. Soc. Ent. de Belg., Ann. 45:375.

Type-species: Crematogaster (Oxygyne) daisyi Forel. Desig. by Wheeler, 1911.
Decacrema Forel, 1910. Soc. Ent. de Belg., Ann. 54:18.

Type-species: Crematogaster schencki Forel. Desig. by Wheeler, 1911.
Crematogaster subg. Atopogyne Forel, 1911. Soc. Vaud. des Sci. Nat., Bui. 47:342.

Type-species: Crematogaster (Atopogyne) hellanica Forel. Desig. by Wheeler, 1911.
Physocrema Forel, 1912. Soc. Ent. de Belg., Mem. 14:220.

Type-species: Crematogaster inflatu^ Smith. Desig. by Wheeler, 1911.
Xiphocrema Forel, 1913. Zool. Jahrb., Abt. f. System. 36:80.

Type-species: Crematogaster tetracantha Emery. Desig. by Emery, 1922.
Eu,crema. Santschi, 1918. Soc. Ent. de France, Bui., p. 182.

Type-species: Formica acuta Fabricius. Orig. desig.
Nematocrema Santschi, 1918. Soc. Ent. de France, Bui. p. 182.

Type-sp>ecies: Crematogaster stadelmanni Mayr. Orig. desig.
Neocrema Santschi, 1918. Soc. Ent. de France, Bui., p. 182.

Type-sp)ecies: Crematogaster distans Mayr. Orig. desig.
Orthocrema Santschi, 1918. Soc. Ent. de France, Bui. p. 182.

Type-species: Myrmica sordidula Nylander. Orig. desig.
Paracrema Santschi, 1918. Soc. Ent. de France, Bui. p. 182.

Type-species: Crematogaster spengeli Forel. Orig. desig.
Sphaerocrema Santschi, 1918. Soc. Ent. de France, BuL p. 182.

Type-species: Crematogaster kneri Mayr. Orig. desig.
Crematogaster subg. Rhachiocrema Mann, 1919. Harvard Univ., Mus. Comp. Zool., BuL
63:318.

Type-sptecies: Crematogaster (Rhachiocrema) wheeleri Mann. Orig. desig.
Tranopeltoides Wheeler, 1922. Amer. Mus. Novitates 48:10.

Type-species: Tranopelta huberi Forel. Orig. desig.
Crematogaster subg. Colobocrema Wheeler, 1927. Quart. Rev. Biol. 2:31.

Type-species: Crematogaster (Colobocrema) cylindriceps Wheeler. Monotypic
Mesocrema Santschi, 1928. Soc. Ent. de Belg., Bui. 68:33.

Type-species: Crematogaster rasoherini Forel. Desig. by Donisthorpe, 1943.
Apterocrema Wheeler, 1936. Psyche 43:45.




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Type-species: Apterocrema atilla^ii Wheeler. Monotypic.

These ants nest in moderately large colonies under objects in the soil, in wood, in cavities of
plants, in insect galls, in carton nests of their own making, or even in the woodwork of buildings.
Most are omnivorous but show a preference for sweets and some tend honeydew excreting in-
sects. Some are well known house-infesting forms; others have been reported to kill young birds
or gnaw rubber insulation from telephone wires. North American species of this genus had been
separated into two subgenera, subgenus Orthocrema for arizonensis and minutissima and its
subspecies and subgenus Crematogaster for the remaining species. Subgenera are not recog-
nized here.

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20:989-996. —Emery, 1895. Zool. Jahrb.,
Abt. f. System. 8:280-288. —Wheeler, 1919. Psyche 26:111. — Creighton, 1939. Psyche
46:137-140 (subg. Orthocretna). — Buren, 1968. Ga. Ent. Soc, Jour. 3:91-121 (subg.
Crema togas ter).

Taxonomy: Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:258-261 (larvae). —Buren,
1958. N. Y. Ent. Soc, Jour. 66:119-134 (also biological notes). —Brown, 1973. In Meggers,
et al., Tropical Forest Ecosystems in Afr. and S. Amer., pp. 161-185 (generic syn.).
—Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:27-30 (larvae).

Biology: Soulie, 1962. Insectes Sociaux 9:181-195 (colony foundation and development).

Crematogaster arizonensis Wheeler
S. Ariz. Ecology: Apparently arboreal; found nesting in mistletoe on
oaks.
Crematogaster arizonensis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:482. ♂.

Biology: Wheeler, 1912. N. Y. Ent. Soc., Jour. 20:130-133 (also description of female, male).

Crematogaster ashmeadi Mayr
Va. to Fla., w. to Tex. Ecology: Strictly arboreal; nests in twigs and branches.
Crematogaster Ashmeadi Mayr, 1886. Zool.-Bot. CJesell. Wien, Verb. 36:463. 5, cJ.
Crematogaster (Acrocoelia) ashmeadi var. mxitura Wheeler, 1932. N. Y. Ent. Soc., Jour.
40:8. ♂.

Taxonomy: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:8.

Biology: Smith, 1924. Ent. News 35:79. —Cole, 1940. Amer. Midland Nat. 24:46. —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326:28-29 (economic importance). — Leuthold, 1968.
Psyche 75:233-248 (tibial gland scent-trail and trail-laying behavior). —Leuthold, 1968.
Psyche 75:334-350 (recruitment to food).

Crematogaster atkinsoni Wheeler
N. C. s. to Fla., w. to Miss. Ecology: Common in coastal salt-grass marshes

where they often build large carton nests on sedges or bushes. Nests are sometimes a

foot and a half or more above the ground and range in size from the diameter of an egg

to that of a human head.
Crematogaster atkinsoni Wheeler, 1919. Psyche 26:108. ♂.
Crematogaster atkinsoni var. helveola Wheeler, 1919. Psyche 26:109. 9 (9, cJ misdet.).

The female and male are C. ashmeadi.

Biology: Atkinson, 1887. Amer. Nat. 21:770-771. —Smith, 1930. Fla. Ent. 14:4-5. —Wheeler,
1932. N. Y. Ent. Soc., Jour. 40:8-9.

Crematogaster browni Buren
W. Tex., N. Mex., s. Ariz. Ecology: In mountains, usually over 5(X)0 ft.; nests
have been found under rocks.
Creynatogaster brmuni Buren, 1968. Ga. Ent. Soc., Jour. 3:100. ♂, ♀.
califomica Emery. S. Calif.; Mexico. Ecology: Nests in soil in desert or semidesert habitats;
found at bases and on roots of various plants cultivating aphids and coccids. Males are
unusuaUy large for this genus.
Crematogaster lineolata laeviuscula var. califomica Emery, 1895. ZooL Jahrb., Abt. f.
System. 8:285. ♀.

Taxonomy: Wheeler, 1934. Pan-Pacific Ent. 10:135-136.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:70. —Wheeler and Wheeler, 1973. Ants
of Deep Canyon, pp. 87-88.




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Crematogaster cerasi (Fitch)
Que. s. to Ga., w. to S. Dak., Ark., N. Mex. Ecology: Nests under rocks and logs;
sometimes does minor damage when nesting in woodwork and timbers in houses.
Myrtnica cerasi Fitch, 1855 (1854). N. Y. State Agr. Soc., Trans. 14:835. ♀.
Cretnatogaster (Acrocoelia) kennedyi Wheeler, 1930. Psyche 37:58. ♀, ♂.
Crematogaster (Acrocoelia) lineolata cerasi var. punctinodis Enzmann, 1946. N. Y. Ent.
Soc, Jour. 54:91, 93, 96. 5, 9, d.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:282-283. —Wheeler, 1933. Psyche
40:83-84. —Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:250-252 (larva, as
lineolata). —Wheeler and Wheeler, 1973. Ga Ent. Soc., Jour. 8:28 (larva).

Biology: Gaige, 1914. Mich. Univ., Mus. Zool., Occas. Papers 5:8-9. — Headley, 1943. Ohio
Jour. Sci. 43:25. —Morris, 1943. Ind. Acad. Sci., Proc. 52:208. —Wheeler and Wheeler, 1944.
N. Dak. Hist. Quart. 11:245. — KannowsW, 1959. Insectes Sociaux 6:125. — Kulman, 1965.
Jour. Econ. Ent. 58:865 (in cocoons of Thyridopteryx ephemeraeformis). — Smith, 1965. U.
S. Dept. Agr., Tech. Bui. 1326:30-31 (economic importance).

Crematogaster clara Mayr
N. J. s. to Fla., w. to Ind., Mo., Tex. Ecology: Nests in a variety of semi-arboreal
situations in or near swamps, salt marshes, rivers, and streams.
Oecodoma (Atta) bicolar Buckley, 1867. Ent. Soc. Phila., Proc. 6:350. ♂. Preocc. by Smith,

1860.
Crematogaster clara Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:990, 993. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:481 (each caste). —Smith, 1924.
Ent. News 35:80. —Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:255-256 (larva,
as laeviuscula).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:283. —Smith, 1965. U. S. Dept. Agr., Tech.
Bui. 1326:31-32 (economic imjwrtance).

Morphology: Pasteels, Crews, and Blum, 1970. Acad. Soc. Paris, Compt. Rend., Ser. D
271:835-838 (histology of gland secreting the trail pheromone).

Crematogaster coarctata Mayr
Oreg.(?), Nev., Calif. Ecology: Nests under rocks and boulders. Suspected of
now being a rather rare species in coastal Calif, due to displacement by the Argentine
ant.
Crematogaster coarctata Mayr, 1870. Zool.-Bot. (Jesell. Wien, Verh. 20:990, 992. ♀.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:482. — Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bui. 104:207-208.

Biology: MaUis, 1941. South. Calif. Acad. Sci., Bui. 40:70-71. — Essig, 1926. Ins. of West. N.
Amer., p. 859.

Crematogaster colei Buren
W. Tex., N. Mex., Ariz. Ecology: Nests under rocks in grassland situations at
elevations over 5000 ft.
Crematogaster colei Buren, 1968. Ga. Ent. Soc., Jour. 3:108. 9,9.

Crematogaster dentinodis Forel
Ariz.; Mexico. Ecology: Nests under stones in open grasslands and chaparral,
or grass and mesquite, or among shrubs in succulent desert areas.
Crematogaster opaca var. dentinodis Forel, 1901. Soc. Ent. de Belg., Ann. 45:130. 9

Crematogaster depilis Wheeler
Tex., N. Mex., Ariz., Nev., s. Calif.; Mexico. Ecology: Nests in and among
roots and lower stems of various plants throughout desert and semi-desert regions.
Crematogaster lineolata opaca var. depilis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.
24:478. ♂.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. ZooL, BuL 104:209 (worker, not
female). —Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:28-30 (larva).

Biology: Cole, 1966. Brigham Yovmg Univ., Sci. Bui., BioL Ser. 7:16. —Wheeler and Wheeler,
1973. Ants of Deep Canyon, p. 88.

Crematogaster emeryana Creighton
W. Tex., Colo., N. Mex., Utah, Ariz.; Mexico. Ecology: Usually in

mountains at elevations of over 6000 ft.; often constructs carton structures under rocks.
Crematogaster lineolata lineolata var. Emery, 1895. Zool. Jahrb., Abt. f. System. 8:282. ♀.
9, (5. Variety described but not named.




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Crematogaster {Acrocoelia) lineolata emeryana Creighton, 1950. Harvard Univ., Mus.
Comp. ZooL, Bui. 104:213-214. ♂.

Biology: Gregg, 1963. Ants of Colo., pp. 358-360.

Crematogaster hespera Buren
W. Tex., N. Mex., Utah, Ariz., Calif.; Mexico. Ecology: Usually arboreal in
Cottonwood trees along rivers and streams; also in other trees and rotten logs.
Crematogaster hespera Buren, 1968. Ga. Ent. Soc., Jour. 3:98. ♂, ♀.

Crematogaster isolata Buren
W. Tex., N. Mex., s. Ariz. Ecology: Arboreal in oaks in mountain ranges at
altitudes over 5000 ft.
Crematogaster isolata Buren, 1968. Ga. Ent. Soc., Jour. 3:106. ♀.

Crematogaster laeviuscula Mayr
La., Tex., Okla.; Mexico. Ecology: Incipient colonies are often found in twigs
and oak galls, larger colonies in hollow trees and logs. No authentic records east of
Mississippi River.
Oecodoma (Atta) arbarea Buckley, 1867. Ent. Soc. Phila., Proc. 6:349-350. 9,9. Preocc. by

Smith, 1858.
Crematogaster laeviuscula Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:990, 993. ♀.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:481 (each caste). —Cole, 1940.
Amer. Midland Nat. 24:46.

Crematogaster larreae Buren
W. Tex., Ariz., Calif.; Mexico. Ecology: Nests in lower stems and among roots

of the creosote bush. Host: Larrea divaricata.
Crematogaster larreae Buren, 1968. Ga. Ent. Soc., Jour. 3:117. 9, 9, S.

Crematogaster lineolata (Say)
Que., Ont. s. to Fla., w. to N. Dak., Colo., Tex. Ecology: Nests in groimd imder

objects or in logs or stiunps; infests houses and often nests within buildings.
Myrmica lineolata Say, 1836. Boston Jour. Nat. Hist. 1:290. 9, 9, 6.
Myrmica (Morurmarium. (.')) marylandica Buckley, 1867. Ent. Soc. Phila., Proc. 6:339. ♀.

Syn. uncertain.
Myrmica (Monomarium (.')) Columbiana Buckley, 1867. Ent. Soc. Phila., Proc. 6:340. 9,9.

Syn. uncertain.
Crematogaster lineolata lineolata var. lutescens Emery, 1895. Zool. Jahrb., Abt. f. System.

8:282. ♀.
Crematogaster (Acrocoelia) lineolata cerasi var. wheldeni Enzmann, 1946. N. Y. Ent. Soc.,

Jour. 54:92-93, 96. ♀.

Taxonomy: Snaith, 1918. Ent. News 29:19. —Buren, 1944. Iowa State Col., Jour. Sci. 18:288.
— Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:252 (larva as lineolata subopaca).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:1-18. —Wheeler, 1917. Conn. Stote
Geol. and Nat. Hist. Survey Bui. 22:585. — Wesson and Wesson, 1940. Amer. Midland Nat.
24:93. —Wallace, 1945. Conn. State Ent., 44th Rpt., Bui. 488:389 (economics). —Wheeler
and Wheeler, 1963. Ants of N. Dak, pp. 132-133. —Gregg, 1963. Ants of Colo., pp. 357, 359.
— Ayre, 1963. Canad. Ent. 95:712-715 (feeding habits). —Smith, 1965. U. S. Dept. Agr.,
Tech. Bui. 1326:32-33 (economic importance). —Ayre, 1968. Canad. Ent. 100:165-172 (prey
finding, capture, transport). — Ayre, 1969. Canad. Ent. 101:118-128 (trail formation and
group foraging).

Crematogaster marioni Buren
S. Calif.; Mexico. Ecology: Arboreal sp>ecies in manzanita and live oaks.
Crematogaster marioni Buren, 1968. Ga. Ent. Soc., Jour. 3:105. ♀.

Crematogaster minutissima
***authority mismatch
minutissima Mayr. N. C. s. to Fla., w. to Tex. Ecology: Nests have been found in
soil at bases of stumps.
Crematogaster minutissima Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:991, 995. 9,9.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:484. — AVheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:14 (larva).

Biology: Smith, 1928. Ent. News 39:277.

Morphology: Pasteels, Crewe, and Blum, 1970. Paris Acad. Soc., Compt. Rend., Ser. D
271:835-838 (histology of gland secreting the trail pheromone).




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Crematogaster minutissima
***authority mismatch
missouriensis Emery. Nebr., Mo., Tex., Colo., N. Mex.

Creniatogaster victima missuriensis (!) Emery, 1895. Zool. Jahrb., Abt. f. System. 8:287.

Taxonomy: Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:260-261 (larva).

Biology: Gregg, 1963. Ants of Colo., pp. 363-364.

Crematogaster minutissima
***authority mismatch
smithi Creighton. Ariz. (Huachuca Mtns.).

Creniatogaster (Orthocrema) minutissima thomcica Creighton, 1939. Psyche 46:138. ♂.

Preocc. by Santschi, 1921.
Creniatogaster {Orthocrema) minutissima smithi Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bui. 104:205. N. name.

Crematogaster mormonum Emery
Utah, Idaho, e. Oreg. (?), Nev., Calif.; Mexico. Ecology: Most common in
the Great Basin; nests under rocks.
Creniatogaster lineolata coarctata var. mormonum Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:284. ♀.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:482 (each caste).

Biology: Cole, 1942. Amer. Midland Nat. 28:363. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, pp. 88-89.

Crematogaster mutans Buren
Nev., Calif. Ecology: The only North American species of this genus suspected
of being parasitic.
Creniatogaster mutans Buren, 1968. Ga. Ent. Soc., Jour. 3:115. ♂, ♀.

Crematogaster navajoa Buren
N. Mex., Ariz, in the Colorado Plateau region. Ecology: Found under rocks and
at bases of various plants.
Creniatogaster navajoa Buren, 1968. Ga. Ent. Soc., Jour. 3:102. ♂, ♀.
noctuma Buren. S. Utah, n. Ariz.

Creniatogaster noctuma Buren, 1968. Ga. Ent. Soc., Joiu". 3:112. 9, 6, possible 5.

Crematogaster opaca Mayr
Ariz. (Tumacacori Mts., under stones among oaks, 3900 ft.); Mexico. Only a single
Arizona record for the U. S.
Creniatogaster opaca Mayr, 1870. Zool.-Bot. (Jesell. Wien, Verh. 20:989. ♀.

Crematogaster opuntiae Buren
N. Mex., s. Ariz. Ecology: Foimd principally at bases of and foraging on
Opuntia fulgida but also at bases of various other plants.
Creniatogaster opuntiae Buren, 1968. Ga. Ent. Soc., Jour. 3:120. ♂.

Crematogaster pilosa Emery
N. J. s. to Ga. Ecology: Often nests in logs and fallen branches in marshy
situations.
Creniatogaster lineolata pilosa Emery, 1895. Zool. Jahrb., Abt. f. System. 8:285. ♂.
Creniatogaster lineolata subpilosa Wheeler, 1913. Psyche 20:115. Syn. uncertain. Nomen

nudum.
Creniatogaster (Acrocoelia) creightoni Wheeler, 1933. Psyche 40:86. ♀.

Taxonomy: Wheeler, 1933. Psyche 40:85.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:379. —Wheeler, 1906. Amer. Mus.
Nat. Hist., Bui. 22:6-7.

Crematogaster punctulata Emery
Va. s. to Fla., w. to Colo., Ariz.; Mexico. Ecology: Abundant in southern
Great Plains region where they nest in the ground and tend aphids and coccids.
Creniatogaster punctulata Emery, 1895. Zool. Jahrb., Abt. f. System. 8:287. ♀.
Creniatogaster lineolata lineolata var. subopaca Emery, 1895. Zool. Jahrb., Abt. f. System.

8:283. 5, 9, cJ.
Creniatogaster (Acrocoelia) opaca var. texana Santschi, 1929. Wien Ent. Ztg. 46:91. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:479-480.

Biology: Smith, 1927. Ent. News 38:310. —Gregg, 1963. Ants of Colo., pp. 361-362, 364.

Crematogaster rifelna Buren
S. Tex.;Mexico. Ecology: Arboreal, found nesting in live oaks and other trees.
Creniatogaster rifelna Buren, 1968. Ga. Ent. Soc., Jour. 3:96. 5, 9, 6.




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Crematogaster vermiculata Emery
N. C. s. to Fla., w. to Ark., La. Ecology: Arboreal, found only in cypress
swamps.
Crematogaster vermiculata Emery, 1895. Zool. Jahrb., Abt. f. System. 8:286. 5 . Types
mislabeled, not found in Calif, or other western states.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:13 (larva).

Tribe SOLENOPSIDINI
Genus MONOMORIUM Mayr

Monomorium Mayr, 1855. Zool.-Bot. Gesell. Wien, Verb. 5:452.

Type-species: MoTiomorium minutum Mayr. Monotypic.
Phacota Roger, 1862. Berlin. Ent. Ztschr. 6:260. Uncertain syn.

Type-species: Phacota sicheli Roger. Monotypic.
Trichomyrmex Mayr, 1865. Novara Exped., Zool., Formicidae, pt. 2, v. 1, p. 19.

Type-species: Trichxymyrmex rogeri Mayr. Monotypic.
Lampromyrmex Mayr, 1868. Beitr. Naturk. Preuss. 1:92.

Type-species: Lampromyrmex gracillim,us Mayr. Monotypic.
Holcomyrmex Mayr, 1878. Zool.-Bot. Gesell. Wien, Verb. 28:671.

Type-species: Holocomyrmex scabriceps Mayr. Desig. by Bingham, 1903.
Epoecus Emery, 1892. Soc. Ent. de France, Bui. 61: CCLXXVI.

Type-species: Epoecus pergandei Emery. Monotypic.
Wheeleria Forel, 1905. Soc. Ent. de Belg., Ann. 49:171. Preocc. by Tutt, 1895.

Type-species: Wheeleria santschii Forel. Monotypic.
Wheeleriella Forel, 1907. Inst. Sci. Revue 4:145. N. name for Wheeleria Forel.
Epixemis Emery, 1908. Deut. Ent. Ztschr., p. 556.

Type-species: Epixenus andrei Emery. Desig. by Wheeler, 1911.
Monomorium subg. Xeromyrmex Emery, 1915. Ent. Soc. de France, Bui. p. 190.

Type-species: Formica salomonis Linnaeus. Orig. desig.
Monomorium, subg. Parholcom.yrmex Emery, 1915. Ent. Soc. de France, Bui. p. 190.

Type-species: Myrmica gracillima Smith. Orig. desig.
Monomorium. subg. Paraholcomyrmex Emery, 1915. Soc. Ent. de France, Bui. p. 191.

Misspelling ?.
Mitara Emery, 1913. Soc. Ent. de Belg., Ann. 57:261.

Type-sp)ecies: Monomorium laeve Mayr. Orig. desig.
Monomorium subg. Corynomyrmex Viehmeyer, 1916. Arch. f. Naturgesch. 18:134.

Type-species: Monomorium (Corynomyrmex) hospitum Viehmeyer. Monotypic.
Monomorium subg. Isolcomyrmex Santschi, 1917. Soc. Cient. Argentina, An. 84:296.

Type-species: Holcomyrmex santschii Forel. Orig. desig.
Paraphxicota Santschi, 1919. Soc. Ent. de France, Bui., p. 91.

Type-species: Phacota noualhieri Emery. Orig. desig.
Monomorium subg. Equ^strimessor Santschi, 1919. Soc. Ent. de France, Bui. p. 92.

Type-species: Holcomyrmex chobauti Emery. Desig. by Donisthorpe, 1943.
Xenhyboma Santschi, 1919. Soc. Espan. de Hist. Nat., Bol. 19:405. Uncertain syn.

Type-species: Xenhyboma mystes Santschi. Monotypic.
Equessimessor Santschi, 1936. Bui. Soc. Sci. Nat. Maroc. 16:32. Emend.
Isholcomyrmex Santschi, 1936. BuL Soc. Sci. Nat. Maroc. 16:32. Variant spelling.
Ireneidris Donisthorpe, 1943. Ent. Monthly Mag. 79:81.

Typ)e-S{)ecies: Ireneidris myops Donisthorpe. Monotypic.
Monomorium subg. Pharaophanes Bernard, 1952. Inst. Franc. d'Afr. Noire, Mem. 19:238.

Nomen nudum.
Xenoaphaenogaster Baroni Urbani, 1964. Atti della Accad. Gioenia di Sci. Nat. Catania
16:50. Uncertain syn.

Type-species: Xenoaphaenogaster inquilina Baroni Urbani. Orig. desig.

Members of this genus are small in size and are particularly adaptable with respect to nesting
sites. They may nest in various preformed cavities or in soil. Most species are Old World, but
because of their adaptability and size, they are especially susceptible to distribution by com-




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merce. Three of the North American species have been introduced and are common tramp spe-
cies and house pests.

Taxonomy. Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:121-122 (larvae).
— Ettershank, 1966. Austral. Jour. Zool. 14:82-93 (generic syn., list of world species).
— Brown, 1973. In Meggers, et al., Tropical Forest Ecosystems in Afr. and S. Amer., pp.
161-185 (generic syn.). —Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:30-31 (larvae).

Morphology: Blum, 1966. Roy. Ent. Soc. London, Proc. (A) 41: 155-160 (source and specificity

of trail pheromones).

Monomorium destructor (Jerdon)
Tenn., Fla.; throughout tropical regions of world. Ecology: A pantropical
tramp and common house-infesting ant that may nest in soil or in buildings. They are
omnivorous and may feed on various household foods. They have been reported to gnaw
holes in fabrics, rubber goods, remove rubber insulation from electric or telephone
wires, and damage polyethylene cable. Introduced, probably originating from Africa or
the Oriental region.

Atta Destructor Jerdon, 1851. Madras Jour. Lit. and Sci. 17:105. ♂.

Myrmica vastator Smith, 1857. Linn. Soc. London, Jour., Zool. 2:71.

Myrmica basalis Smith, 1858. Cat. Hym. Brit. Mus. 6:125. ♂.

Taxonomy: Bingham, 1903. Faima British India 2:209 (each caste). —Emery, 1908. Deut. Ent.
Ztschr. pp. 665-666, 671. —Phillips, 1934. (Hawaii Univ.) Expt. Sta., Pineapple Prod. Coop.
Assoc, Bui. 15:2. —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:64.

Biology: Wheeler, 1910. Ants, pp. 10, 153, 221. —Wheeler, 1914. Amer. Jour. Trop. Dis. and
Prev. Med. 2:160-168 (economics). —Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:833.
— Kalshoven, 1937. Ent. Meded. van Nederland. Indie 3:65-71. — Marlatt, 1928. U. S. Dept.
Agr., Farmers' Bui. 740:9. — Kempf, 1960. Studia Ent. 3:506-507 (economics). —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326:37-38 (economic importance).

Monomorium floricola (Jerdon)
Fla., Ala.; throughout tropical regions of world. Ecology: An arboreal

species, nesting in twigs and branches or under bark of trees or other plants. Commonly
infests houses and feeds on household foods. Introduced, probably originating from
tropical Asia.
Atta floricola Jerdon, 1851. Madras Jour. Lit. and Sci. 17:107. ♂.
Monomarium speculare Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. Kl., Sitzber. 53:509.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:88 (female). —Smith, 1936. Puerto
Rico Univ., Jour. Agr. 20:831, 834. —Wheeler and Wheeler, 1955. Amer. Midland Nat.
54:121 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:64-65.

Biology: Wheeler, 1910. Ants, pp. 153, 426. —Smith, 1930. Fla. Ent. 14:3. —Wheeler, 1932. N.
Y. Ent. Soc., Jour. 40:9. —Plank and Smith, 1940. Puerto Rico Univ., Jour. Agr. 24:49-75
(association with pineapple mealybug, Psevdococcus brevipes (Ckll.)). — Brown, 1964. Ent.
News 75:15. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:36-37 (economic importance).

Monomorium minimum (Buckley)
Que., Ont. s. to Fla., w. to Mont., Colo., Calif.; Mexico. Rare or absent in

the Pacific Northwest. Ecology: Nests in exposed soil, under cover of objects, or in

rotting or faulty wood. Sometimes invades houses from outdoors or nests in woodwork

Parasite: Monomorium pergandei (Emery). Little black ant.
Myrmica (Monomarium (.')) minima Buckley, 1867. Ent. Soc. Phila, Proc. 6:338. ♂, ♀.
Myrmica (Monomarium (.')) atra Buckley, 1867. Ent. Soc. Phila., Proc. 6:342. 9 (?).
Monomorium minutum ergatogyna Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:269. ♂.

apterous ergatoid 9 .
Monomorium minimum emersoni Gregg, 1945. Psyche 52:66. ♂, ♀.
Monomorium metoecus Brown and Wilson, 1957. Ent. News 68: 239-244. Ergatogyne.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:274-275 (each caste). —Wheeler,
1905. South. Calif. Acad. Sci., Bui. 40:60. —Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.
24:423. — Buren, 1944. Iowa State Col., Jour. Sci. 18:289. —Wheeler and Wheeler, 1955.
Amer. Midland Nat. 54:122 (larva). —Wilson and Brown, 1958. Ent. News 69:33-38 (worker
of metoecus; also biology). — Ettershank, 1966. Austral Jour. Zool. 14:90 (syn. of
metoecus).




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Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:274, 377. — Marlatt, 1922. U. S. Dept.
Agr., Farmers' Bui. 740:4, 10. — Eckert and MaUis, 1937. Calif. Agr. Expt. Sta., Cir. 342:23.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31:271, 272, 274, 279-280. — Metcalf and Flint, 1939.
Destructive and Useful Insects, p. 770. —Gregg, 1944. Ent. Soc. Amer., Ann. 37:454,456,
466. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 133-135. —Gregg, 1963. Ants of
Colo., pp. 366-368. —Bums, 1964. Ent. Soc. Amer., Ann. 57:138 (association with tuliptree
scale). —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:33-34 (economic importance).
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 89-90.

Monomorium pergandei (Emery)
D. C. Ecology: Supposedly workerless and parasitic. The host colony

contained only winged males and winged females. Host: Monomorium minimum

(Buckley).
Epoecus pergandei Emery, 1892. Soc. Ent. de France, Bui. 61: CCLXXVI. 9,6.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:272-274. —Wheeler, 1910. Ants, p.

498. —Smith, 1947. Amer. Midland Nat. 37:569 (female). — Ettershank, 1966. Austral. Jour.

Zool. 14:82, 91.

Monomorium pharaonis
***authority mismatch
(Linnaeus). Fla., throughout U. S. and Canada in larger cities; cosmopolitan.

Ecology: Not uniformly distributed; adapted to field conditions only in Florida but
probably found in every town or city of commercial importance especially in hotels,
large apartment buildings, groceries, or other places where food is commercially
handled. Colonies are populous and are commonly found in the most inaccessible places
in buildings. They breed continuously through the year and take 38 to 42 days to
develop. They may feed on various household foods, damage silk, rayon and rubber
goods and are also known to damage insect collections. This is the most persistent and
difficult of all our house-infesting ants to control or eradicate. Introduced, probably
originating from Africa or tropical Asia. Pharaoh ant.

Formica Pharaonis Linnaeus, 1758. Syst. Nat., Ed. 10:580.

Taxonomy: Emery, 1908. Deut. Ent. Ztschr., pp. 664-665, 667, 684 (each caste).

— Donisthorpe, 1927. British Ants, p. 104. — Buren, 1944. Iowa State Col, Jour. Sci. 18:289.
—Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:121 (larva). —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62: 16 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog.
14:68-60.

Biology: BeUevoye, 1889. U. S. Dept. Agr., Insect Life 2:230-233 (in 2 years, 1,360,000
workers, 1,900 females, and 560 males taken on same premises). — Herrick, 1914. Insects
Injurious to the Household and Annoying to Man, pp. 174-176. — Essig, 1926. Insects of
West. N. Amer., p. 857. —Metcalf and Flint, 1939. Destructive and Useful Insects, p. 770.
—Smith, 1934. Kans. Acad. Sci., Trans. 37:140-142. — Armand, 1942. Pests 10:18-19
(thermal preference as guide to control work). — Peacock and Baxter, 1949. Ent. Monthly
Mag. 85:256. —Peacock, 1950. Ent. Monthly Mag. 86:129. —Peacock and Baxter, 1950. Ent.
Monthly Mag. 86:171-178 (life history). —Peacock, 1950. Ent. Monthly Mag. 86:294.
—Peacock, et al., 1950. Scot. Dept. Agr., Misc. Pub. 17:1-50 (biology and control).
—Peacock, 1951. Ent. Monthly Mag. 87:185-191. —Sudd, 1953. Advancement of Sci. (Brit.)
10:17-18 (colony foundation). —Peacock, et al., 1954. Ent. Monthly Mag. 90:154-158 (male
production by parthenogenesis). —Peacock, et al., 1955. Ent. Monthly Mag. 91:125, 130.
—Peacock, et al., 1955. Ent. Monthly Mag. 91:37-42 (viability in regard to temperature
and moisture). —Brown, 1958. Acta Hym. 1:36. —Wheeler and Wheeler, 1963. Ants of N.
Dak, pp. 135-136. —Brown, 1964. Ent. News 75:14-15. — Eichler, 1963. Deut. Ent. Ztschr.
10:207-215. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:34-36 (economic importance).
— Wisniewski, Myjak, and Przyborowsld, 1971. Polskie Pismo Ent. 61:459-474 (biology, in
Poland). —Petersen and Buschinger, 1971. Anz. f. Schadlingsk. u. Pflanzensch. 44:121-127
(behavior). —Petersen and Buschinger, 1971. Ztschr. f. Angew. Ent. 68:168-175 (behavior
of sexuals).

Morphology: Tanquary, 1913. 111. State Lab. Nat. Hist., Bui. 9:443-453 (physiology). —Hall
and Smith, 1951. Ent. Monthly Mag. 87:217: — HaU and Smith, 1952. Ent. Monthly Mag.
88:97-102 (thoracic structure). —Hall and Smith, 1953. Evolution 7:127. —HaU and Smith,
1954. Ent. Monthly Mag. 90:176-182 (somatic mosaics). —Smith and Peacock, 1957. Roy.
Soc. Edinb., Proc., Section B, 66:235-261 (cytology).




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Monomorium viridum
***authority mismatch
peninsulatum Gregg. Ga., Fla. w. to Colo., Ariz., s. Calif. Ecology: Most commonly
found in soil under objects.
Monomorium peninsulatum Gregg, 1945. Psyche 52:62. ♂, ♀.

Biology: Gregg, 1963. Ants of Colo., pp. 368-370.

Monomorium viridum
***authority mismatch
viridum Brown. N. Y. (Coram, Selden, L. I.), N. J. (Lakehurst). Ecology: The types
from N. J. were taken from nests with 5 to 8 inch craters in yellow and white sand
among scanty, low weeds of a roadside strip.
Monomorium viridum. Brown, 1943. Ent. News 54:243. 9,9.

Genus XENOMYRMEX Forel

Xenomyrmex Forel, 1885. Soc. Vaud. des Sci.Nat., Bui. 20:369.

Type-species: Xenomyrmex stollii Forel. Monotypic.
Myrm,ecineUa Wheeler, 1922. Amer. Mus. Novitates 46:1.

Type-species: Myrmecinella panamana Wheeler. Monotypic.

A small New World genus of three species. The ants are arboreal, nesting in small colonies in
plant cavities such as twigs, galls, and thorns.

Revision: Wheeler, 1931. Rev. de Ent. 1:129-139. — Creighton, 1957. Amer. Mus. Novitates
1843:1-14.

Taxonomy: Creighton, 1953. Amer. Mus. Novitates 1634:2. —Wheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:19-20 (larva). — Ettershank, 1966. Austral. Jour. Zool. 14:150-152.

Xenomyrmex floridanus
***authority mismatch
floridanus Emery. Fla.; Bahamas, Cuba. Another subspecies is found in Mexico.
Xenomyrmex stolli floridanus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:275. 5, d.
Xenomyrmex stolli floridanus var. lucayamis Wheeler, 1905. Amer. Mus. Nat. Hist., Bui.

21:87. ♂.
Xenomyrmex stolli rufescens Wheeler, 1931. Rev. de Ent. 1:133,137. ♀.
Xenomyrmex stolli cubanu^s Wheeler, 1931. Rev. de Ent. 1:134-135. ♀.

Biology: Wheeler, 1901. Amer. Nat. 35:538. —Wheeler, 1932. N. Y. Ent. See., Jour. 40:6.

Morphology: Holldobler, 1971. Jour. Ins. Physiol. 17:1497-1499 (sex pheromone).

Genus SOLENOPSIS Westwood

Solenopsis Westwood, 1841. Ann. and Mag. Nat. Hist. 6:86.

Type-species: Soletuypsis mandibularis Westwood. Monotypic.
Diplorhoptrum Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:449.

Type-species: Formica fugax Latreille. Monotypic.
Solenopsis subg. Synsolenopsis Forel, 1918. Soc. Vaud. des Sci. Nat., Bui. 52:155.

Type-species: Solenxypsis bruchiella Emery. Monotypic.
Solenopsis subg. Diagyne Santschi, 1923. Rev. Suisse de Zool. 30:267.

Type-sjjecies: Solenopsis succinea Emery. Monotypic.
Labauchena Santschi, 1930. Soc. Ent. Argent., Rev. 13:81.

Type-sjjecies: Labattchena daguerrei Santschi. Monotypic.
Solenopsis subg. Euophthalma Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66: 43.

Type-sp»ecies: Myrmica globtilaria Smith. Orig. desig.
Solenopsis subg. Oedaleocerus Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66:43.

Type-species: Solenopsis argulata Emery. Orig. desig.
Bisolenopsis Kusnezov, 1953. Acta. Zool. Lilloana [Tucuman] 13:1.

Type-species: Bisolenopsis sea Kusnezov. Monotypic.
Paranamyrma Kusnezov, 1954. Mus. Entre Rios, Mem. 30:9.

Type-species: Pamnamyrmxi solenopsidis Kusnezov. Monotypic.
Lilidris Kusnezov, 1957. Zool. Anz. 158:268, 274. Uncertain syn.

Type-species: Lilidris metatarsalis Kusnezov. Monotypic.
Solenopsis subg. Granisolenopsis Kusnezov, 1957. Zool. Anz. 158:270, 277.

Type-species: Soletwpsis (Granisolenopsis) granivora Kusnezov. Monotypic.




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Previously, this genus was divided into several subgenera, the subgenus Solenopsis included
the larger, polymorphic species aurea, geminata, invicta, richteri, and xyloni, the subgenus
Euophthalma included globularia littoralis and huachuca7ia, and the subgenus Diplorhcrptruyn
included the minute "thief ants", the remaining species listed here. Those previously placed in
the subgenus Solenopsis are commonly known as "fire ants", the colonies are populous, nests are
usually mounded in exposed situations, and workers are agressive and possess a painful sting.
They are of the most economically important of ants. Most species in the other subgenera are
minute and cryptic, often lestobiotic in the nests of other ants.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:277-279 {Diplorhoptrum). —Wheeler,
1915. Amer. Mus. Nat. Hist., Bui. 34:397. — Creighton, 1930. Amer. Acad. Arts and Sci.,
Proc. 66:39-139.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:131-136 (larvae). — Snelling,
1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3, 10 pp. (keys to all castes of subg.
Solenopsis). — Ettershank, 1966. Austral. Jour. Zool. 14:134-144 (generic syn. and list of
world species). — Buren, 1972. Ga. Ent. Soc, Jour. 7:1-27 (revisionary studies of the
imported fire ants).

Morphology: Blum, et al., 1958. Science 128:306-307 (chemical, insecticidal and antibiotic
properties of fire ant venom). — MacConnel, Blum and Fales, 1971. Tetrahedron
26:1129-1139 (chemistry of fire ant venom). —San Martin, 1971. In Bucheil and Buckley,
Venomous Animals and Their Venoms v. 3, pp. 95-101 (venomous ants of Solenopsis).

Solenopsis aurea Wheeler
Tex., N. Mex., Ariz., s. Calif.; Mexico. Ecology: Nests are in fully exposed
situations, in dry, coarse, gravelly soil, and are without a mound.
Solenopsis geminata var. aurea Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:336. 5,2,

6.
Solenopsis aurea amblychila Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:394. 5, 9, S.

Taxonomy: Snelling, 1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3: 7.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:425-426.

Solenopsis carolinensis Forel
Mass. s. to N. C, Tenn., Ohio. Ecology: Probably lestobiotic.

Solenopsis tenana race carolinensis Forel, 1901. Soc. Ent. de Belg., Ann. 45:345. 5 , 9 , 6.

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:92.

Solenopsis geminata (Fabricius)
S. C. s. to Fla., w. to Tex. and s. to Peru; West Indies; tropical Asia and
Pacific Islands. Ecology: In the U. S., it is most common on or near the coast with its
incidence decreasing inland except in Florida where it is distributed over most of the
state. Nests are in the ground in open areas in dry to moist soil of various composition
surmounted by irregular piles of dirt; they may also be under cover of objects or in
rotting logs. Economic importance is similar to that of xyloni McCook, but its
importance has subsided since the introduction of the imported fire ants. A common
tramp species, native to the New World and spread by commerce to other tropical
regions of the world. Fire ant.

Atla geminata Fabricius, 1804. Systema Piezatorum, p. 423. ♀.

Atta Rufa Jerdon, 1851. Madras Jour. Lit. and Sci. 17:106. ♀.

Solenopsis mandifmlaris Westwood, 1841. Ann. and Mag. Nat. Hist. 6:87. ♂.

Myrmica virulens Smith, 1858. Cat. Hym. Brit. Mus. 6:132. ♂.

Atta clypeata Smith, 1858. Cat. Hym. Brit. Mus. 6:169. ♀, ♂.

Solenopsis cephalotes Smith, 1858. Linn. Soc. London, Jour., Zool. 3:149. ♀.

Crematogaster laboriosus Smith, 1860. Linn. Soc. London, Jour., Zool. 4:109. ♀.

Diplorhoptrum drewseni Mayr, 1861. Eur. Formicid., p. 71. ♂.

Myrmica glaber Smith, 1862. Ent. Soc. London, Trans. (3) 1: 34. ♂.

Myrmica polita Smith, 1862. Ent. Soc. London, Trans. (3) 1: 34. ♀.

Myrmica {Monomarium (.')) saxicola Buckley, 1867. Ent. Soc. Phila, Proc. 6:341. 9. Syn.
uncertain.

Atta Lincecumii Buckley, 1867. Ent. Soc. Phila., Proc. 6:344. 5 , %. Syn. uncertain.

Atta brazoensis Buckley, 1867. Ent. Soc. Phila., Proc. 6:345. 5 , %. Syn. uncertain.

Atta colaradensis Buckley, 1867. Ent. Soc. Phila., Proc. 6:346. 5, 'H. Syn. uncertain.




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Solenopsis geminata var. diabola Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:424. 5 , 'U,
S.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:132-133 (larva). — Snelling,
1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3:7-9. —Wilson and Taylor, 1967.
Pacific Ins. Monog. 14:58-59.

Biology: Mann, 1920. Amer. Mus. Nat. Hist., Bui. 42:427. — Marlatt, 1928. U. S. Dept. Agr.,
Farmers' Bui. 740:5. — Neig, 1930. Bombay Nat. Hist. Soc, Jour. 34:185. —Clark, 1931.
Tex. Agr. Expt. Sta. Bui. 435:1-12. —Cole, 1934. Ent. Soc. Amer., Ann. 27:395. — PhiUips,
1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc, Bui. 15:12-17. —Smith, 1936.
Puerto Rico Univ., Jour. Agr. 20:838-839. —Plank and Smith, 1940. Puerto Rico Univ.,
Jour. Agr. 24:49-76. —Travis, 1941. Fla. Ent. 24:15-22. —Griffiths, 1942. Science 96:271-272.
— Lindquist, 1942. Jour. Econ. Ent. 35:850-851. — Kempf, 1961. Studia Ent. 4:507. —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326:40-41 (economic importance). —Banks, Plumley, and
Hicks, 1973. Ent. Soc. Amer., Ann. 66:234-235 (polygyny in a colony). —Bass and Hays,
1976. Ga. Ent. Soc, Jour. 11: 34-36 (in S. C).
globularia littoralis Creighton. N. C. s. to Fla., w. to La.; Mexico. Ecology: Commonly found

on open beaches; nests are constructed in or vmder rotten logs. S. globularia globularia

(Smith) occurs in Central and S. Amer.
Solenopsis (.Euophthalma) globularia littoralis Creighton, 1930. Amer. Acad. Arts and

Sci., Proc 66:110, 113. 5, 9, 6.
Solenopsis globularia mobilensis Smith, 1931. Ent. News 42:20. 5 . Nomen nudum.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:22 (larva).
Biology: Smith, 1931. Ent. News 42:20. —Smith, 1933. Fla. Ent. 17:23.

Solenopsis huachucana Wheeler
Ariz. (Miller Canyon, Huachuca Mtns.).

Solenopsis huachucana Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:393. ♂, ♀.

Taxonomy: Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66:118-120.

Solenopsis invicta Buren
N. C, S. C, Ga., Fla., Tenn., Ala., Miss., Ark., La., Tex.; Brazil. Ecology: The
most widespread of the two imported fire ants and the most economically important
being an annoying and sometimes destructful pest. The mound nests are unsightly and
sometimes interfere with agricultural operations; workers are agressive and can inflict a
painful sting; workers steal seeds from seedbeds and feed on germinating seeds of com,
may gnaw holes in various fabrics, foster honeydew excreting insects, injure or kill
young rabbits, pigs, and other mammals, quail and other birds, gnaw into roots, stems,
buds, and fruits of various agricultural crops, and may girdle young plants. Introduced,
probably originating from Mato Grosso, Brazil; the earliest U. S. record is 1945 from
Daphne, Ala. Red imported fire ant. Because of the recent distinction of two introduced
species in the U. S., it is difficult to determine whether much of the literature pertains
to this species or S. richteri. Only selected references are given below; for a
bibliography through 1971 see U. S. Dept. Agr., Coop. Econ. Ins. Rpt., 1971, Sept., 21
(36): 639-652. Much of the literature is under the name S. saevissima richteri Forel.
Solenopsis invica (!) Buren, 1972. Ga. Ent. Soc, Jour. 7:9-15. 5, 9, 6.

Taxonomy: Wilson, 1951. Evolution 5:68-79. —Wilson, 1952. Inst. Oswaldo Cruz, Mem.
50:49-68. —Wilson, 1953. Evolution 7:262-263. —Brown, 1957. Quart. Rev. Biol. 32:258-261.
— Wilson and Brown, 1958. Evolution 12:211-218 (morphological changes in introduced
population). —Snelling, 1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3:10. —Buren,
1972. Ga- Ent. Soc, Jour. 7:9-15. —Buren, et. al., 1974. N. Y. Ent. Soc, Jour. 82: 113-124
(zoogeography of the imported fire ants).

Biology: WUson, 1958. Sci. Amer. 198:36-41. —Bellinger, Dyer, King and Pratt, 1965. Ga.
Acad. Sci., Bui. 23:122 (review of fu-e ant problem). — Fincher and Lund, 1967. Ga. Ent.
Soc, Jour. 2:91-94 (biology and life cycle in Ga.). — Rhoades and Davis, 1967. Jour. Econ.
Ent. 60:544-558 (effects of meteorological factors on biology and control). — Anon. 1971
Sept., U. S. Dept. Agr., Coop. Econ. Ins. Rpt., 21 (36): 639-652 (bibliography through 1971).
—Collins and Markin, 1971. Ent. Soc Amer., Ann. 64:1376-1380 (inquilines and other
arthropods).— Markin and Dillier, 1971. Ent. Soc. Amer., Ann. 64:562-565 (seasonal life




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cycle in Miss.). —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:41^3 (economic
importance). —Wilson and Oliver, 1969. Jour. Econ. Ent. 62:1268-1271 (food habits).
— Glancey, et at, 1970. Nature 226:863-864 (pheromone may induce brood tending).
—Wilson, Dillier, and Markin, 1971. Ent. Soc. Amer., Ann. 64:660-665 (foraging territories).
— Markin, et al., 1971. Ga. Ent. Soc., Jour. 6:145-156 (nuptial flight and flight ranges).
—Collins and Markin, 1971. Ent. Soc. Amer., Ann. 64:1376-1380 (inquilines and other
arthropods in nests). —Markin, Collins, and Dillier, 1972. Ent. Soc. Amer., Ann.
65:1053-1058 (colony founding by queens). —Harris and Bums, 1972. Environ. Ent.
1:362-365 (predation on lone star tick). — Bhatkar, et al, 1972. Environ. Ent. 1:274-279
(confrontation behavior between Lasius neoniger and the imported fire ant). — Stringer, et
al., 1972. Jour. Econ. Ent. 65:872-873 (air separation of different castes). —Glancey,
Stringer, and Bishop, 1973. Ga. Ent. Soc., Jour. 8:217-220 (trophic egg production).
—Glancey, et al., 1973. Ga. Ent Soc., Jour. 8:237-238 (multiple fertile queens in colonies).
—O'Neal and Markin, 1973. Ga. Ent. Soc., Jour. 8:294-303 (brood nutrition and parental
relationships). —Hubbard, 1974. Ga. Ent. Soc, Jour. 9: 127-132 (influence of nest material
and colony odor on digging). —Allen, et. al., 1974. Ent. Soc. Amer., Ann. 67: 43^6
(distribution and habits in Brazil). —Morrill, 1974. Environ. Ent. 3: 265-271 (production and
flight). —Morrill, 1975. Ga. Ent. Soc., Jour. 10: 162-164 (reduction of populations by tillage).
— Horton, et al., 1975. Ga. Ent. Soc., Jour. 10: 207-213 (food carrying ability and
recruitment time). — Lofgren, Banks, and Glancey, 1975. Ann. Rev. Ent. 20: 1-30 (biology
and control of imported fire ants). —Bass and Hays, 1976. Ga. Ent. Soc, Jour. 11: 34-36 (in
S. C). —Adams, et al., 1976. Ga. Ent. Soc, Jour. 11: 165-169 (economic importance, impact
on soybean harvest).

Morphology: Blum, et al., 1958. Science 128:306-307 (chemical, insecticidal, and antibiotic
properties of venom). —Callahan, Blum, and Walker, 1959. Ent. Soc Amer., Ann.
52:573-590 (histology of poison glands and sting). — Adrouny, Derbes, and Jung, 1959.
Science 130:479 (hemolytic component of fire ant venom). — Wilson, Diu-lach and Roth,
1959. Psyche 65:108-114 (chemical releases of necrophoric behavior). — Wilson, 1959.
Science 129:643-644 (nature of odor trail). —Wilson, 1962. Animal Behavior 19:134-147
(chemical communication among workers-mass foraging). — Blum and Callahan, 1960. XI
Int. Kong. Ent. Wien, Verb. B. 111:290-293 (chemical and biological properties of venom).
— Wilson, 1962. Animal Behavior 10:148-158 (chemical communication among workers).
— Wilson, 1962. Animal Behavior 10:159-164 (induction of social responses). — Walker and
Clower, 1961. Ent. Soc. Amer., Ann. 54:92-99 (alimentary canal of queen). — Caro, Derbes,
and Jung, 1957. Amer. Med. Assoc, Arch. Dermat. 75:475^488 (skin responses to sting).
—Thompson and Blum, 1967. Ent. Soc. Amer., Ann. 60:632-642 (spermatozoa). —Vinson,
1970. Ent. Soc. Amer., Ann. 63:930-935 (gustalory response to various electrolytes).
— MacConnell, Blum, and Fales, 1970. Science 168:840-841 (alkaloid from fire ant venom).
—Smith and Smith, 1971. Arch. Dermat. 103:438-441 (multiple fire ant stings - a
complication of alcoholism). — MacConnell, Blum, and Fales, 1971. Tetrahedron
26:1129-1139 (chemistry of fire ant venom). —Glancey, et al., 1976. Ga. Ent. Soc, Jour. 11:
83-88 (testes degeneration). — Robeau and Vinson, 1976. Ga. Ent. Soc, Jour. 11: 198-203
(effects of juvenile hormone analogues on caste differentiation).

Solenopsis krockowi Wheeler
N. Mex.; Mexico.

Solenopsis krockcrwi Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:428. ♂, ♀.

Solenopsis molesta (Say)
Ont. s. to Fla., w. to Wash., Calif. Ecology: Lestobiotic, usually nests in or near
nests of other ants from which they rob food and brood. A house-infesting ant and of
great annoyance because of their small size; sometimes nests in woodwork and masonry
of houses. Thief ant. The form occurring west of the Rockies is sometimes referred to
as the subspecies validvuscula.
Myrmica molesta Say, 1836. Boston Jour. Nat. Hist. 1:293. ♀.
Myrmica minuta Say, 1836. Boston Jour. Nat. Hist. 1:294. ♂.
Myrmica (Tetmamxrrium (.')) exigtia Buckley, 1867. Ent. Soc. Phila., Proc. 6:342. 5, "9" =

6.
Solenopsis debilis Mayr, 1886. ZooL-Bot. Gesell. Wien, Verb. 36:461. 5, 9, 6.
Solenopsis molesta var. validiuscula Emery, 1895. Zool. Jahrb., Abt. f. System. 8:278. ♀.




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Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:430. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bui. 22:584 (each caste). — Wheeler and Wheeler, 1955. Amer.
Midland Nat. 54:134-135 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:377-378. — McColloch and Hayes,
1916. Jour. Econ. Ent. 9:23-38. —Hayes, 1920. Kans. Agr. Expt. Sta., Tech. Bui. 7:1-54.
— Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342:21-22. — Metcalf and Flint, 1939.
Destructive and Useful Insects, p. 770. —Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:72.
— Macnamara, 1945. Canad. Ent. 77:40. — Grundmann and Peterson, 1953. Kans. Ent. Soc,
Jour. 26:59. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 136-138. —Gregg, 1963.
Ants of Colo., pp. 372-375. — Ayre, 1963. Canad. Ent. 95:712-715 (feeding habits). —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326:43^5 (economic importance). — Wheeler and
Wheeler, 1973. Ants of Deep Canyon, pp. 91-92.

Solenopsis pergandei Forel
Va. s. to Fla., w. to La. Ecology: Nests have been found in soil in rotting
stumps, and next to nests of other ants.
Solenopsis pergandei Forel, 1901. Soc. Ent. de Belg., Ann. 45:343. $, 9, 6.

Taxonomy: Smith, 1931. Ent. News 42:20. —Wheeler and Wheeler, 1960. Ent. Soc. Wash.,
Proc. 62:21-22 (larva).

Biology: Smith, 1931. Ent. News 42:20. —Smith, 1944. Fla. Ent. 27:15.

Solenopsis picta Emery
S. C. s. to Fla., w. to Tex. Ecology: Nests have been found in hollow twigs.
Solenopsis tenuis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:262. ♂. Preocc. by Mayr,

1877.
Solenopsis picta Emery, 1895. Zool. Jahrb., Abt. f. System. 8:278. ♀.
Solenopsis picta var. moerens Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:393. ♂.

Taxonomy: Smith, 1942. Ent. Soc. Wash., Proc. 44:211. —Wheeler and Wheeler, 1960. Ent.
Soc. Wash., Proc. 62:22 (larva).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:10.

Solenopsis pilosula Wheeler
Tex. (Alice).

Solenopsis pilosula Wheeler, 1908. Am6r. Mus. Nat. Hist., Bui. 24:426. 5 , d .

Solenopsis richteri Forel
Miss., Ala.; S. Brazil, Uruguay, Argentina. Ecology: A mound building ant with
similar habits and importance of invicta Buren. May have been more widespread in U.
S. until the introduction of invicta which may have replaced richteH in many localities.
Introduced, probably originating from southern S. Amer.; first recorded from Mobile,
Ala. in 1930 but may have been present there for 10 to 12 years. Black imported fire
ant. Until 1972, the imported fire ant had been known under one name, S. saevissima

Solenopsis richteri Forel
; consequently, it is difficult to determine to which species, either invicta
or richteri, much of the earlier literature pertains. For a bibliography of the "imported
fire ant" through 1971, see U. S. Dept. Agr., Coop. Econ. Ins. Rpt., 1971, Sept., 21 (36):
639-652, as well as the list of references given under invicta.
Solenopsis Pylades var. Richteri Forel, 1909. Deut. Ent. Ztschr., p. 267. ♂, ♀.

Taxonomy: Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66:87 (first U. S. record).
—Wilson, 1951. Evolution 5:68-79. — WUson, 1952. Inst. Oswaldo Cruz, Mem. 50:49-68.
—Wilson, 1953. Evolution 7:262-263 (origin of variation). —Brown, 1957. Quart. Rev. Biol.
32:258-261. —Wilson and Brown, 1958. Evolution 12:211-218 (morphological changes in
introduced population). — Buren, 1972. Ga. Ent. Soc., Jour. 7:4-8 (two species of imported
fire ants).

Biology: Green, 1952. Jour. Econ. Ent. 45:593-597 (biology and control in Miss.). — Green,
1967. Miss. State Univ., Agr. Expt. Sta., Bui. 737, 23 pp.

Solenopsis salina Wheeler
W. Tex., Colo. w. to CaUf.; Mexico. Ecology: Nests under rocks and wood;
lestobiotic.
Solenopsis salina Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:427. ♂.

Biology: Gregg, 1963. Ants of Colo., pp. 376-378.

Solenopsis tennesseensis Smith
Fla., Tenn., Miss., Tex., Kans., Calif, (nr. Azusa).

Solenopsis (Diplorhoptrum) longiceps Smith, 1942. Ent. Soc. Wash., Proc. 44:210. ♀.
Preocc. by Forel, 1907.




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Solenopsis (Diplorhoptrum) tennesseensis Smith, 1951. In Muesebeck, et al., U. S. Dept.
Agr., Agr. Monog. 2:814. N. name.

Solenopsis texana
***authority mismatch
catalinae Wheeler. Calif. (Catalina Is.).

Solenopsis texana catalinae Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:269. ♂, ♀.

Taxonomy: Wheeler, 1905. South. Calif. Acad. Sci., Bui. 4:60.

Solenopsis texana
***authority mismatch
texana Emery. Ont. s. to Fla., w. to Okla., Tex. Ecology: Nests have been found in logs,
stumps, and under bark.
Solenopsis pollux var. texana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:278. ♂.
Solenopsis rosella Kennedy, 1938. Canad. Ent. 70:232. 5, 9, 6.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:430-431. —Wheeler and Wheeler,
1955. Amer. Midland Nat. 54:136 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14:70.

Solenopsis truncorum Forel
N. C. s. to Fla., w. to Colo., Ariz., Calif. Ecology: Nests have been found
under rocks and other objects; lestobiotic.
Solenopsis texana race truncorum Forel, 1901. Soc. Ent. de Belg., Ann. 45:346. 2, 9.
Solenopsis molesta var. castariea Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:430. 2 .

Biology: Gregg, 1963. Ants of Colo., pp. 378-380.

Solenopsis xyloni McCook
N. C. s. to Fla., w. to Colo., Calif.; Mexico. Ecology: Nests are in ground in
exposed soil or under cover of stones or other objects, sometimes in wood; they are
surmounted by irregular and variable-shaped mounds of loose soil. A serious pest in
some parts of its range: builds ugly mounds on lawns, inflicts painful stings, steals seeds
from seedbeds, kills young poultry and other birds, girdles nursery stock, gnaws into
buds, tubers, and fruits of various plants, bites holes in fabrics such as silk, nylon, and
linen, removes rubber insulation from telephone wires, and feeds on household foods.
Southern fire ant.

Solenopsis xyloni McCook, 1879. In Comstock, Rpt. Cotton Ins., p. 188. ♂, ♀.

Myrmica (Atta) sabeana Buckley, 1867. Ent. Soc. Phila., Proc. 6:343. 5 . Syn. questionable.

Solenopsis geminata maniosa Wheeler, 1915. Amer. Mus. Nat Hist., Bui. 34:396. ^ ,%, 9 ,
6.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:133 (larva). — Snelling, 1963.
Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3:9.

Biology: Severin, 1923. Jour. Econ. Ent. 16:96-97. —Smith, 1936. Jour. Econ. Ent. 29:120-122.
— Eckert and MaUis, 1937. Calif. Agr. Expt. Sta. Cir. 342:10-21. — Mallis, 1938. Pan-Pacific
Ent. 14:87-91. — Metcalf and Flint, 1939. Destructive and Useful Insects, p. 771.
— Eagleson, 1940. Jour. Econ. Ent. 33:700. — Wray, 1962. Jour. Econ. Ent. 55:145 (apparent
introduction into N. C). —Gregg, 1963. Ants of Colo., pp. 370-371, 373. —Smith, 1965. U. S.
Dept. Agr., Tech. Bui. 1326:38-40 (economic importance). —Wheeler and Wheeler, 1973.
Ants of Deep Canyon, pp. 92-94.

Morphology: Blum, Roberts, and Novak, 1961. Psyche 68:73-74 (chemical and biological
characters of venom).

Genus OLIGOMYRMEX Mayr

Oligomyrmex Mayr, 1867. Ent. Tidskr. 10:110.

Type-sp)ecies: Oligomyrmex concinnus Mayr. Monotypic.
Aeromyrma Forel, 1891. Soc. Ent. de Belg., Ann. 35:307.

Type-species: Aeromyrma nosindam.bo Forel. Monotypic.
Pheidologeton subg. Aneleus Emery, 1900. Termes. Fuzetek 23:327.

Type-species: Solenopsis sim.ilis Mayr. Desig. by Wheeler, 1911.
Erebomyrma Wheeler, 1903. Biol. Bui. 4:138.

Type-sp>ecies: Erebomyrma longii Wheeler. Monotypic.
Pheidologeton subg. Lecanomyrma Forel, 1913. Soc. Vaud. des Sci. Nat., Bui. 49:56.

Type-species: Pheidologeton (Lecanom,yrma) butteli Forel. Monotypic.
Oligomyrmex subg. Octella Forel, 1915. Arkiv for Zool. 9:69.

Type-species: Oligomyrmex {Octella) pachycerus Forel. Monotypic.




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Spelaeomyrmex Wheeler, 1922. Amer. Mus. Novitates 45:9.

Type-species: Spelaeomyrmex urichi Wheeler. Monotypic.
Oligom,yrmex subg. Hendecatella Wheeler, 1927. Lab. Zool. Gen. e Agr. Portici, Bol. 20:93.

Type-species: OUgomyrmex (Hendecatella) capreolus Wheeler. Monotypic.
Solenopsis subg. Solenops Karawajew, 1930. Zool. Anz. 92:207. Preocc. by Dufour, 1820.

Type-species: Solenopsis (Solenops) weyeri Karawajew. Monotypic.
Sporocleptes Arnold, 1948. Natl. Mus. South. Rhodesia, Occas. Papers 2 (14): 219.

Type-species: Sporocleptes nicotiana Arnold. Monotypic.
Solenopsis subg. Crateropsis Patrizi, 1948. 1st. Ent. Univ. Bologna, Bol. 17:174.

Type-sp»ecies: Solenopsis (Crateropsis) elmenteitae Patrizi. Monotypic.
Nimbamyrma Bernard, 1953. Mem. Inst. Franc. Afr. Noire 19, fasc. 1, p. 240. Questionable
syn.

Type-species: Nimbamyrma villiersi Bernard. Monotypic.

A large genus in tropical Africa and Asia, with fewer species in the neotropics and only one
species reaching the United States. Colonies are small and collections are usually made from
under bark of logs, in rotten wood, or leaf litter. Most species may be lestobiotic in the nests of
other ants and termites.

Taxonomy: Wilson, 1962. Psyche 69:62-72. — Ettershank, 1966. Austral. Jour. Zool. 14:119-124
(generic syn. and list of world species). — Wheeler and Wheeler, 1953. Psyche 60:141
(larvae).

Oligomyrmex longii (Wheeler)
Okla. (Ft. Sill), Tex. (Denton). Ecology: May be lestobiotic.
Erebomyrma longii Wheeler, 1903. Biol. Bui. 4:140. 5, 9, S.

Taxonomy: Mann, 1926. Psyche 33:104. —Wheeler, 1936. Amer. Acad. Arts and Sci., Proc.
71:197.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:423. —Wheeler, 1910. Ants, pp. 113,
140, 152, 158-159, 427-428.

Tribe LEPTOTHORACINI

Genus MACROMISCHA Roger

Macromischa Roger, 1863. Berlin. Ent. Ztschr. 7:184.

Type-species: Macromischa purpurata Roger. Desig. by Wheeler, 1911.
Macromischa subg. Croesomyrmex Mann, 1920. Amer. Mus. Nat. Hist., Bui. 42:408.

Type-sjjecies: Macromischa (Croesomyrmex) wheeleri Mann. Orig. desig.
Macromischa subg. Antillaemyrmex Mann, 1920. Amer. Mus. Nat. Hist., Bui. 42:408.

Type-species: Macromischa (Antillaemyrmex) terricola Mann. Orig. desig.

A rather large genus in the neotropics with only three species reaching the United States.

Revision: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:138-142. —Mann, 1920. Amer. Mus.
Nat. Hist., Bui. 42:407^24. —Wheeler, 1931. Harvard Univ., Mus. Comp. Zool., Bui. 72:3-34.
—Wheeler, 1937. Harvard Univ., Mus. Comp. Zool., Bui. 81:441, 449-458, 463^65. —Smith,
1939. Ent. Soc. Amer., Ami. 32:502-509.

Taxonomy: Brown, 1973. In Meggers, et al.. Tropical Forest Ecosystems in Afr. and S.
Amer., p. 181 (listed as a provisional syn. of Leptothorax).

Macromischa floridana (Wheeler)
Fla. (Paradise Key, Dade Co.). Ecology: Nest of the types was found in a
hollow branch of a small tree.
Antillaemyrmex floridanus Wheeler, 1931. Harvard Univ., Mus. Comp. Zool., Bui. 72:27.
5.

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:11.

Macromischa polita Smith
Ariz. Ecology: One nest was found under the bark of a tree, but most specimens
have been taken on the groimd.
Macromischa polita Smith, 1939. Ent. Soc. Amer., Ann. 32:503, 506. ♂.

Macromischa subditiva Wheeler
La., Tex.; Mexico. Ecology: Nests have been found under willow bark, and
from dead hollow branches lying on the ground.
Macromischa subditiva Wheeler, 1903. Psyche 10:99. ♀.




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Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14:73. —Smith, 1939. Ent. Soc.
Amer., Ann. 32:506. — Creighton, 1965. Psyche 72:282.

Genus ROGERIA Emery

Rogeria Emery, 1894. Soc. Ent. Ital., Bol. 26:188.

Type-species: Rogeria curvipubens Emery. Desig. by Wheeler, 1911.
Rogeria subg. Irogeria Emery, 1915. Soc. Ent. de France, Bui. p. 191.

Type-species: Rogeria procera Emery. Orig. desig.

A neotropical genus with about 24 species. Only two species reach the southwestern U. S.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:28 (larvae). — Kempf, 1961.

Rev. Brasil. Biol. 21:435-141. —Kempf, 1962. Studia Ent. 5:1-38. —Kempf, 1963. Rev.

Brasil. Biol. 23:189-196 (S. Amer. species). —Kempf, 1964. Studia Ent. 7:45-71. —Kempf,

1965. Rev. Brasil. Biol. 25:181-186. — Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci.

No. 236:1-6 (first record of genus in U. S.). —Wheeler and Wheeler, 1973. Psyche 80: 74

(larvae, revised description).

Rogeria creightoni Snelling
Tex. (La Feria, Cameron Co.).

Rogeria creightoni Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. No. 236:2-4. ♂.

Rogeria huachucana Snelling
Ariz. (Cochise Co.). Ecology: Specimens found beneath stones.

Rogeria huachucana Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. No. 236:4-6. ♂.

Genus LEPTOTHORAX Mayr

Other than those genera listed under Macromischa and the subgenera of Leptothorax, Brown
(1973) gives the following synonyms or possible synonyms: Chalepoxenus Menozzi, 1923?,
Doronomyrmex Kutter, 1945?, Epimyrma Emery, 1915?, Formicoxenus Mayr, 1855 ?, Gonepi-
myrma Bernard, 1948 ?, Icothorax Hamann and Klemm, 1967 ?, Leoytomyrma Amoldi, 1968 ?,
Myrmamnophilus Menozzi, 1924, Myrmetaerus Soudek, 1925 ?, Myrmoxenus Ruzksy, 1902 ?,
Symmyrmica Wheeler, 1904 ?, and Temnothorax Mayr, 1861. There is apparently considerably
more work to be done on this and related genera; consequently, I am using subgenera in Lep-
tothorax as has been done in past catalogs.

Taxonomy: Brown, 1973. In Meggers, et al.. Tropical Forest Ecosystems in Afr. and S.
Amer., pp. 161-185.

Genus LEPTOTHORAX Subgenus NESOMYRMEX Wheeler

Leptothorax subg. Goniothorax Emery, 1896. Soc. Ent. Ital., Bol. 28:26, 58. Preocc. by
Mibie-Edwards, 1879.

Type-sp)ecies: Leptothorax xncinus Mayr. Desig. by Wheeler, 1911.
Nesomyrmex Wheeler, 1910. Amer. Mus. Nat. Hist., Bui. 28:259.

Type-species: Nesomyrmex clavipilis Wheeler. Monotypic.
Leptothorax subg. Caulomyrma Forel, 1914. Soc. Vaud. des Sci. Nat., Bui. 50:233.

Type-species: Leptothorax echinatinodis Forel. Orig. desig.
Limnomyrmex Arnold, 1948. Nat. Mus. So. Rhodesia, Occas. Papers 2 (14): 222.

Type-species: Limnomyrmex stramineus Arnold. Monotypic.

Most sp)ecies of this subgenus are neotropical

Revision: Kempf, 1959. Studia Ent. 2:391-432.

Taxonomy: Smith, 1950. Psyche 57:30. —Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann.
48:25 (larva). —Brown, 1971. Breviora 365:4-5 (generic syn.).

Leptothorax wilda Smith
S. Tex.; Mexico. Ecology: Arboreal, forms small colonies in plant cavities;
probably nocturnal.
Leptothorax {Goniothorax) urilda Smith, 1943. Ent. Soc. Wash., Proc. 45:155. 9,9.

Biology: Creighton, 1971. Ga. Ent. Soc., Jour. 6:207-210 (distribution and habits).

Genus LEPTOTHORAX Subgenus MYRAFANT Smith

Leptothorax subg. Myrafant Smith, 1950. Psyche 57:29.




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Type-species: Leptothorax curvispinosus Mayr. Orig. desig.

These small ants nest in small colonies, commonly in preformed cavities under bark, hollow
twigs, dried grass stems, old galls, or empty nut shells. A few may be found under rocks or in
the soil. Some species are lestobiotic, nesting near the nests of other ants, and others may be du-
lotic.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317-318, 320-323. —Wheeler, 1903.
Acad. Nat. Sci. Phila., Proc. 55:215, 224, 232-256.

Taxonomy: Smith, 1950. Psyche 57:29. —Smith, 1952. N. Y. Ent. Soc., Jour. 60:96-106
(tricaririatus - texanus complex). —Brown, 1955. Ent. News 66:43. — Wheeler and
Wheeler, 1955. Ent. Soc. Amer., Ann. 48:22-25 (larvae). —Cole, 1956. Tenn. Acad. Sci., Jour.
31:30-31 (tricarinatus - texanics complex). — Cole, 1958. Ent. Soc. Amer., Ann. 51:535-538
(nitens - carinatus complex). — Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:23
(larvae).

Leptothorax ambiguus
***authority mismatch
ambiguus Emery. Que. to Va., w. to N. Dak., S. Dak., Iowa, Nebr. Ecology: Nests in

soil in woodlands and grasslands or in hollow dead grass stems near the soil.
Leptothorax (Leptothorax) curvispinosus ambiguus Emery, 1895. Zool. Jahrb., Abt. f.

System. 8:317, 320. ♂.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22:588-589.
— Wesson and Wesson, 1940. Amer. Midland Nat. 24:97 (each caste). — Creighton, 1950.
Harvard Univ., Mus. Comp. Zool., Bui. 104:261-262. —Wheeler and Wheeler, 1955. Ent.
Soc. Amer., Ann. 48:22-23 (larva).

Biology: Sturtevant, 1925. Psyche 32:314. — Buren, 1944. Iowa State Col., Jour. Sci. 18:287.
— Kannowski, 1959. Insectes Sociaux 6:124, 150-151, 155 (parasitism; pleometrosis).
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 140-141.
ambig^uus foveatus Smith. 111. (Plainfield). Ecology: Found in nest of a species of
Aphaenogaster in a roadside ditch.
Leptothorax foveata Smith, 1934. Psyche 41:211. ♂.
ambigruus pinetorum Wesson and Wesson. Ohio (Jackson Co.).

Leptothorax ambiguus var. pinetorum Wesson and Wesson, 1940. Amer. Midland Nat.
24:97. 5, 9, 6.

Leptothorax andrei Emery
N. Mex., Ariz., Nev., Calif. Ecology: Nests have been found imder stones.

Leptothorax (Leptothorax) andrei Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 322. ♂.

Taxonomy: Cole, 1958. Ent. Soc. Amer., Ann. 51:537-538 (each caste).

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:73. —Wheeler and Wheeler, 1973. Ants
of Deep Canyon, p. 95.

Leptothorax bradleyi Wheeler
Ga., Ala., Fla. Ecology: Colonies found in galleries in bark.
Leptothorax bradleyi Wheeler, 1913. Psyche 20:113. ♂.

Taxonomy: Wilson, 1952. Ent. News 63:68-71 (each caste).

Leptothorax carinatus Cole
W. Tex., Ariz. Ecology: Nests were found beneath stones.

Leptothorax (Leptothorax) carinatus Cole, 1957. Tenn. Acad. Sci., Jour. 32:213-215. ♂, ♀.

Taxonomy: Cole, 1958. Ent. Soc. Amer., Ann. 51:537-538. —Wheeler and Wheeler, 1973.
Psyche 80: 70-71 (semipupa).

Leptothorax curvispinosus Mayr
Maine s. to Fla., w. to Iowa, Kans., Okla., Tex., Ariz. Ecology: Nests in
plant cavities such as hollow stems, twigs, and in acorns. Enslaved by Leptothorax

Leptothorax duloticus Wesson
and Harpagoxenus americanu^ (Emery).
Leptothorax curvispinosus Mayr, 1866. Akad. der Wien, Math.-Nat. KL, Sitzber. 53:508. ♂.
Stenamma gallarum Patton, 1879. Amer. Nat. 13:126. ♂, ♀.

Taxonomy: Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36:451, 453 (worker, female). —Cole,
1940. Amer. Midland Nat. 24:56-57.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:385. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31:289, 305. — Headley, 1943. Ent. Soc. Amer., Ann. 36:743-753 (population studies).




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—Talbot, 1957. Ecology 38:449-456 (populations). — Kannowski, 1959. Insectes Sociaux
6:124. —Wilson and Pagan, 1974. N. Y. Ent. Soc, Jour. 82: 106-112 (estimation of total
behavioral repertories).

Leptothorax furunculus Wheeler
Wyo., Colo.

Leptothorax furunculus Wheeler, 1909. N. Y. Ent. Soc, Jour. 17:82. ♀.

Biology: Gregg, 1963. Ants of Colo., pp. 381-382.

Leptothorax gallae Smith
Calif. Ecology: Commonly found on the canyon live oak, Quercus chrysolepis,
often in twig galls made by cynipids.
Leptothorax (Leptothorax) gallae Smith, 1949. Psyche 56:112. ♀.

Leptothorax hispidus Cole
W. Tex.; Mexico. Ecology: Nests were found under stones at higher elevations.
Leptothorax (Leptothorax) hispidus Cole, 1957. Tenn. Acad. Sci., Jour. 32:42-45. 2,9.

Taxonomy: Wheeler and Wheeler, 1973. Psyche 80: 71 (immatures).
longrispinosus Roger. Que., Ont. s. to Ga., Ala. w. to Iowa. Ecology: Nests in plant cavities such
as hollow stems, twigs, and in acorns. Enslaved by Leptothorax duloticus Wesson and
Harpagoxenus americayius (Emery).
Leptothorax longispinostts Roger, 1863. Berlin. Ent. Ztschr. 7:180. ♂.
Leptothorax (Leptothorax) longispinosus laeviceps Buren, 1944. Iowa State Col., Jour. Sci.

18:286. ♂.
Leptothorax longispinosus iowensis Buren, 1945. Ent. Soc. Wash., Proc. 47:288. N. name
for laeviceps, thought to be preocc. by leviceps Emery, 1898.

Taxonomy: Wheeler, 1917. Conn. State (Jeol. and Nat. Hist. Survey Bui. 22:588. —Cole, 1940.
Amer. Midland Nat. 24:56-57. —Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:24
(larva).

Biology: Wheeler, 1910. Ants, pp. 212, 222, 495, 504. —Dennis, 1938. Ent. Soc. Amer., Ann.
31:274, 289, 305. — Headley, 1943. Ent. Soc. Amer., Ann. 36:743-753 (population studies).
—Kannowski, 1959. Insectes Sociaux 6:125. — Lettendre and Pilon, 1972. Nat. Canad.
99:73-82 (ecology; in Que.).

Leptothorax mariposa Wheeler
Calif. (Yosemite Natl. Pk.). Ecology: Colonies were found under stones.
Leptothorax nitens var. mariposa Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:507.

Taxonomy: Cole, 1958. Ent. Soc. Amer., Ann. 51:536.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:73-74.

Leptothorax minutissimus Smith
D. C. The type series of several females was associated with L.
curvispinosu^ workers, but the exact relationship is not known.
Leptothorax minutissimus Smith, 1942. Ent. Soc. Wash., Proc. 44:59. ♀.

Leptothorax nevadensis
***authority mismatch
eldoradensis Wheeler. Calif. (Coastal Range and lower elevations of the Sierras).
Leptothorax eldoradensis Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:414. 0.

Biology: Mallis, 1941. South. Calif. Acad. Sci., BuL 40:73.

Leptothorax nevadensis
***authority mismatch
melanderi Wheeler. W. Mont, to e. Wash.

Leptothorax melanderi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:81. ♀.

Leptothorax nevadensis
***authority mismatch
nevadensis Wheeler. Wash., Oreg., Calif. (Eastern slopes of Sierras north to
Cascade Mtns.). Ecology: Nests in soil, usually under stones.
Leptothorax nevadensis Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 252. 5, 9, d.

Taxonomy: Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:81. —Wheeler and Wheeler, 1973. Psyche
80: 71 (larva).

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 95-96 (subspecies not
recognized).

Leptothorax nevadensis
***authority mismatch
rudis Wheeler. Nev., Calif. (Sierras from Lake Tahoe to Sequoia Park).

Leptothorax nevadensis rudis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:508. ♂.
9.
Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:73. —Cole, 1966. Brigham Young Univ.,
Sci. Bui., Biol. Ser. 7:17.




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Leptothorax nitens Emery
Wyo., Colo., Tex. w. to Wash., Oreg., Calif. Ecology: Found under rocks and in
duff.
Leptothorax (Leptotharax) nitens Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 322. ♂.
Leptothorax nitens occidentalis Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:233, 245. ♂.
Leptotharax nitens var. heathii Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 245. ♂.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:369-370. —Cole, 1958. Ent. Soc. Amer., Ann.
51:536. —Wheeler and Wheeler, 1973. Psyche 80: 71, 73 (larva).

Biology: MalUs, 1941. South. Calif. Acad. Sci., Bui. 40:73. —Gregg, 1963. Ants of Colo., pp.
382-384. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 96.

Leptothorax obliquicanthus Cole
Colo., N. Mex. Ecology: Foimd under stones in meadows and prairies.
Leptotharax (Myrafant) obliquicanthus Cole, 1953. Ent. Soc. Wash., Proc. 55:28-30. ♂.

Taxonomy: Gregg, 1953. Breviora 22:1-3.
Biology: Gregg, 1963. Ants of Colo., pp. 384-385, 387.

Leptothorax obturator Wheeler
Tex. Ecology: One colony was found in an oak gall.

Leptothorax obturator Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 249. 5, 9, 6.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:24 (larva).

Biology: Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19:663-664. —Wheeler, 1910. Ants, pp.
208-209.

Leptothorax rugatulus
***authority mismatch
brunnescens Wheeler. N. Dak., Mont. s. to Colo., Utah. Ecology: Nests under rocks
or wood. Sometimes not distinguished from the typical subspecies in the literature.
Leptotharax rv^atulv^ brunnescens Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.

52:510. ♂.
Leptotharax rugatulus dakotensis Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11:247.
5.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:25 (larva).

Biology: Gregg, 1963. Ants of Colo., pp. 388-389.

Leptothorax rugatulus
***authority mismatch
rugatulus Emery. N. Dak., S. Dak., Colo., N. Mex. w. to B. C, Calif. Ecology: Nests

under stones, wood, in decaying wood, in grasses.
Leptotharax {Leptothorax) rugatulus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317, 321.

5.
Leptotharax curvispinosu^ rugatulus var. Cockerelli Wheeler, 1903. Acad. Nat. Sci. Phila.,

Proc. 55:241. ♂, ♀.
Leptothorax curvispinosus annectens Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:242. ♀.
Leptotharax rugatulus var. mediarufus Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.

52:510. ♂, ♀.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 241. —Cole, 1942. Amer.

Midland Nat. 28:369. — Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104:267-269.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:509. —Cole, 1934. Psyche 41:222.
—Cole, 1954. Tenn. Acad. Sci., Jour. 29:240. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 141-143 (does not distinguish subspecies). —Gregg, 1963. Ants of Colo., pp. 386-388.

Leptothorax schaumi Roger
Maine to Ga., w. to Iowa., Kans., Tex. Ecology: Commonly nests in the bark of
trees.
Leptotharax schaumi Roger, 1863. Berlin. Ent. Ztschr. 7:180. ♂.
Leptotharax fartinodis Mayr, 1886 Zool.-Bot. Gesell. Wien, Verb. 36:451. 2,9.
Leptotharax fartinodis var. gilxms Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:235. ♂, ♀.
Leptotharax fartinodis var. melanoticus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:235.
5, 9.

Taxonomy: Wesson and Wesson, 1940. Amer. Midland Nat. 24:94-96. —Cole, 1940. Amer.
Midland Nat. 24:56. —Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104:269-271.
—Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:23 Oarva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:385. —Wheeler, 1916. Ind. Acad. Sci.,
Proc. 26:461.




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Leptothorax schmittii Wheeler
Colo. (Canyon City).

Leptotharax Schmittii Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 242. ♂.

Taxonomy: Gregg, 1963. Ants of Colo., pp. 388, 390-391.

Leptothorax silvestrii (Santschi)
S. Ariz. Ecology: Nests in evergreen oaks at altitudes of 3500 ft. or more.
Tetramorium silvestrii Santschi, 1909. Soc. Ent. Ital., Bol. 41:6. ♂.

Taxonomy: Creighton, 1953. Amer. Mus. Novitates 1635:1-7 (each caste described; note on
habitat).

Leptothorax stenotyle Cole
Ariz. (nr. Rustler's Park, Chiricahua Mtns.). Ecology: Nests were found under
stones.
Leptothorax (Leptotharax) angustinodus Cole, 1956. Tenn. Acad. Sci., Jour. 31:28-30. ♂, ♀.

Preocc. by Stitz, 1917.
Leptothorax (Leptothorax) stenotyle Cole, 1956. Tenn. Acad. Sci., Jour. 31:214. N. name.

Leptothorax terrigena Wheeler
Tex. (Austin).

Leptothorax terrigena Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 254. ♂, ♀.

Leptothorax texanus
***authority mismatch
davisi Wheeler. N. Y., N. J., Fla.

Leptothorax texanus davisi Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:385. ♂, ♀.

Taxonomy: Smith, 1952. N. Y. Ent. Soc., Jour. 60:104-106.

Leptothorax texanus
***authority mismatch
texanus Wheeler. Mich. (?), Ohio, s. to N. C, Ga. w. to Okla., Tex. Ecology: Nests have
been found in sandy soil.
Leptotharax texanus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223-245. 5, 9, 6.

Taxonomy: Gregg, 1944. Ent. Soc. Amer., Ann. 37:446. —Smith, 1952. N. Y. Ent. Soc, Jour.
60:102-104. —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:24 (larva).

Biology: Smith, 1932. Ent. News 43:160. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:98.

Leptothorax tricarinatus
***authority mismatch
neomexicanus Wheeler. W. Colo., Utah, N. Mex., Ariz. Ecology: Nests found in
soil, under stones, in open grassy areas.
Leptotharax neomexicanus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223-248. ♂.

Taxonomy: Smith, 1952. N. Y. Ent. Soc., Jour. 60:100-102.

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22:341. —Cole, 1953. Ent. Soc. Wash.,
Proc. 55:28. —Gregg, 1963. Ants of Colo., pp. 393-394.

Leptothorax tricarinatus
***authority mismatch
tricarinatus Emery. N. Dak, S. Dak., Iowa w. to Wyo., Colo., Utah. Ecology:
Nests under rocks and in soil.
Leptotharax (Leptotharax) tricarinatus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318,
321. ♂.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 247-248. — Buren, 1944. Iowa
State Col., Jour. Sci. 18:286, 288. —Smith, 1952. N. Y. Ent. Soc., Jour. 60:98-100 (worker,
male).

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., p. 143. —Gregg, 1963. Ants of Colo.,
pp. 391-393.

Leptothorax tuscaloosae Wilson
N. C, Ala. Ecology: Colony was found in small cavity in soU under a bed
of moss at base of an oak tree.
Leptotharax (Myrafant) tuscaloosae Wilson, 1950. Psyche 57:128-130. 2,9.

Leptothorax wheeleri Smith
N. C, Ga., Fla., Ohio, Tenn., Ala., Miss. Ecology: Colonies found in cavities in
trees and under bark.
Leptotharax wheeleri Smith, 1929. Ent. Soc. Amer., Ann. 22:547. ♂, ♀.

Taxonomy: Wilson, 1952. Ent. News 60:67-68, 70 (worker, male).

Biology: Smith, 1931. Ent. News 42:18. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:90, 96.

Genus LEPTOTHORAX Subgenus DICHOTHORAX Emery

Leptotharax subg. Dichothorax Emery, 1895. Zool. Jahrb., Abt. f. System. 8:323.




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Type-species: Leptotkorax {Dichothorax) pergandei Emery. Desig. by Wheeler,
1911.

The status of the two forms below needs clarification. Though regarded as subspecies, the
ranges overlap considerably.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 323-324. —Wheeler, 1903. Acad.
Nat. Sci. Phila., Proc. 55:224, 256-260.

Leptothorax pergandei
***authority mismatch
floridanus Emery. N. J., N. C. to Fla., w. to Tex. Ecology: Has been found in
stumps, logs, and nut shells.
Leptotkorax (Dichothorax) floridanus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 324.

Leptotkorax (Dickotkorax) pergajidei flavus Smith, 1929. Ent. Soc. Amer., Ann. 22:549. ♂.

9.
Leptotkorax (Dickotkorax) pergandei floridantis var. spinosus Smith, 1929. Ent. Soc.

Amer., Ann. 22:551. ♂.

Taxonomy: Smith, 1931. Ent. News 42:18-19.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:290, 305.

Leptothorax pergandei
***authority mismatch
pergandei Emery. D. C. s. to Ga., Tenn., w. to Nebr., Tex. Ecology: Apparently a
ground nesting species.
Leptotkorax {Dickotkorax) pergandei Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 323.

5.
Leptotkorax {Dickotkorax) manni Wesson, 1935. Ent. News 46:208. 5 , 9 , 6. Preocc. by
Wheeler, 1914.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 256-259 (worker, female).
—Smith, 1924. Ent. News 35:50. —Cole, 1940. Amer. Midland Nat. 24:56, 58. —Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:23 (larva).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:290. — Buren, 1944. Iowa State Col., Jour.
Sci. 18:288.

Genus LEPTOTHORAX Subgenus LEPTOTKORAX Mayr

Leptotkorax Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:431.

Type-species: Formica acervorum Fabricius. Desig. by Bingham, 1903.
Myckothorax Ruzsky, 1904. Sapiski Imp. Russ. Geog. Obshch. 41:288.

Type-sjjecies: Formica acervorum Fabricius. Orig. desig.

In North America, this subgenus is confined mostly to the northern and western United
States and Canada. The ants nest in living and dead trees, decaying wood, soil, or stumps. Some
are inquilines in nests of other ants and some are dulotic.

Taxonomy: Brovm, 1955. Ent. News 66:43-50. —Wheeler and Wheeler, 1955. Ent. Soc. Amer.,
Ann. 48:21-22 (larvae).

Leptothorax crassipilis Wheeler
Wyo., Colo., N. Mex., Utah, Ariz., Nev. Ecology: Found under rocks, under
wood, in decaying logs.
Leptotkorax (Myckotkomx) acervorum crassipilis Wheeler, 1917. Amer. Acad. Arts and
Sci., Proc. 52:513. 5, 9, cJ.

Taxonomy: Cole, 1954. Tenn. Acad. Sci., Jour. 29:240-241.

Biology: Cole, 1953. Ent. Soc. Wash., Proc. 55:27. —Gregg, 1963. Ants of Colo., pp. 401^02,
405.

Leptothorax diversipilosus Smith
Wash. Ecology: An inquitine in nests of Form,ica obscuripes Forel.
Leptotkorax (Myckotkorax) diversipilosus Smith, 1939. Ent. Soc. Wash., Proc. 41:179. ♀.
ergatoid 9.

Taxonomy: Smith, 1956. Ent. Soc. Wash., Proc. 58:271-275 (female, ergatoid male).

Biology: Alpert and Akre, 1973. Ent. Soc. Amer., Ann. 66:753-760 (distribution, abimdance,
behavior).




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Leptothorax duloticus Wesson
Mich., Ohio, 111. (?). Ecology: Dulotic, enslaves Leptothorax curvispinosus
Mayr and L. longispinosus Roger.
Leptothorax (Mychothorax) duloticus Wesson, 1937. Ent. News 48:125. ♂, ♀.

Taxonomy: Wesson, 1940. Brooklyn Ent. Soc, Bui. 35:81-83 (male, biology).

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:94. — Talbot, 1957. Ecology
38:449-456 (populations). — Kannowski, 1959. Insectes Sociaux 6:124-215, 150-151
(parasitism).
hirticomis Emery. N. Dak., S. Dak., Colo., Utah., Calif. Ecology: An inquiline in nests of
Formica obscuripes Forel and possibly Formica integroides integroides Emery.
Leptothorax (Leptothorax) hirticomis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317, 319.

5.
Leptothorax (Leptothorax) hirticomis formidolostis Wheeler, 1915. Amer. Mus. Nat. Hist.,
Bui. 34:415. «?, ergatoid 9.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223-224. —Smith, 1939. Ent. Soc.
Wash., Proc. 41:176-179 (worker, ergatoid female).

Biology: Weber, 1935. Ecol. Monog. 5:200. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart.
11:248. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 145-146. —Gregg, 1963. Ants of
Colo., pp. 403, 405. — Snelling, 1965. South. Calif. Acad. Sci., Bui. 64:16 (also description of
dealate female and ergatoid male).

Leptothorax muscorum (Nylander)
Newfoimdland (Labrador) w. to N. W. T., Alaska, s. to Conn., Wis., N.

Mex., Ariz., Calif.; Eurasia. Ecology: Has been found at 69° 22' N., Kidluit Bay on

Richards Is., the northernmost record of any New World ant. Mostly found in

woodlands, in decaying stumps, logs, under bark of fallen trees, or under rocks. It is

best able to survive in extreme arctic conditions under stones.
Myrmica muscorum Nylander, 1846. Acta Soc. Sci. Fenn. 2:1054. 5, 9, 6.
Leptothorax canadensis Provancher, 1887. Addit. Corr. Faune Ent. Canada, Hym. p. 245.

5, 9,cJ.
Leptothorax (Leptothorax) canadensis var. yankee Emery, 1895. Zool. Jahrb., Abt. f.

System. 8:317, 319. ♂, ♀.
Leptothorax yankee var. kincaidi Pergande, 1900. Wash. Acad. Sci., Proc. 2:520. 9,9.
Leptothorax muscorum var. sordidv^ Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223,

224. ♂.
Leptothorax acervorum canadensis var. convivialis Wheeler, 1903. Acad. Nat. Sci. Phila.,

Proc. 55:228. 9,9.
Leptothorax canadensis obscurus Viereck, 1903. Amer. Ent. Soc., Trans. 29:72. ♀.
Leptothorax (Leptothorax) acervorum canadensis var. Calderoni Forel, 1914. Deut. Ent.

Ztschr., p. 617. 9, 9.
Leptothorax (Mychothorax) muscorum var. septentrionalis Wheeler, 1917. Amer. Acad.

Arts and Sci., Proc. 52:511. 9, 9, d.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 225-229. —Wheeler, 1917.
Conn. State Geol. and Nat. Hist. Survey Bui. 22:588. —Wheeler, 1917. Amer. Acad. Arts
and Sci., Proc. 52:512-513. —Brown, 1955. Ent. News 66:47-50 (also ecology). —Wheeler
and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:21 flarva). — Francoeur and Beique, 1966.
Canad. Ent. 98:142 (Provancher ant types).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:621. —Wheeler and Wheeler, 1944. N.
Dak. Hist. Quart. 11:247-248. — Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:73. —Cole,
1942. Amer. Midland Nat. 28:369-370. —Gregg, 1946. Amer. Midland Nat. 35:748.
—Kannowski, 1959. Insectes Sociaux 6:125. —Wheeler and Wheeler, 1963. Ants of N. Dak,
pp. 143-145. —Gregg, 1963. Ants of Colo., pp. 393-400. —Chapman, 1969. Ent. Soc. Amer.,
Ann. 62:1256-1259 (release and recovery of marked winged forms). — Moglich, Machwdtz,
and HoUdobler, 1974. Science 186: 1046-1047 (tandem calling).




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Leptothorax provancheri Emery
Que., Maine w. to Alta., N. Dak., Colo., N. Mex. Ecology: Apparently an

inquiline, found in nests of Myrmica incompleta incompleta Provancher and M.

lobicomis fracticomis Emery.
Leptothorax (Leptothorax) provancheri Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317,

320. ♂.
Leptothorax emersoni Wheeler, 1901. Amer. Nat. 35:433. $ , 9 , cJ .
Leptothorax emersoni glacialis Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. (n. s.) 5:71. ♂, ♀.

Leptothorax (Mychothorax) emersoni hirtipilis Wheeler, 1917. Amer. Acad. Arts and Sci.,

Proc. 52:515. ♀.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 229. —Cole, 1954. Tenn. Acad.
Sci., Jour. 29:241. —Wheeler and Wheeler, 1973. Psyche 80: 73-74 (larva).

Biology: Wheeler, 1907. Wis. Nat. Hist. Soc., Bui. (n. s.) 5:78-83. —Wheeler, 1910. Ants, pp.
107, 393, 434-436. —Wheeler, 1901. Amer. Nat. 35:436-438. —Wheeler, 1903. Jour. Psychol,
and Neurol. 2:1-21. — Kannowski, 1957. PsycRe 64:1-5. —Wheeler and Wheeler, 1963. Ants
of N. Dak., pp. 146-148. —Gregg, 1963. Ants of Colo., pp. 404-407.

Genus SYMMYRMICA Wheeler

Symmyrmica Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:1, 3.

Type-species: Symmyrmica chamberlini Wheeler. Monotypic.

Taxonomy: Brown, 1973. In Meggers ei al.. Tropical Forest Ecosystems in Afr. and S.
Amer., p. 185 (possible syn. of Leptothorax).

Symmyrmica chamberlini Wheeler
Utah, Oreg. Ecology: An inquiline in nest of Manica m,utica (Emery).
Symmyrmica chamberlini Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:5. 5,9, apterous
ergatoid 6.

Biology: Wheeler, 1910. Ants, pp. 432-434. —Wheeler, 1919. Amer. Phil. Soc., Proc. 58:22.

Genus HARPAGOXENUS Forel

Tomognathus Mayr, 1861. Die Europaischen Formiciden, pp. 29, 56. Preocc. by Dixon,
1850.

Type-sf>ecies: Myrmica sublaeiris Nylander. Monotypic.
Harpagoxenus Forel, 1893. Soc. Ent. de Belg., Ann. 37:167. N. name for Tomognathus.
Protomognathus Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:3.

Type-sp)ecies: Tomognathus americanus Emery. Monotypic.

Ants of this genus are dulotic, enslaving and conducting raids on colonies of certain species of
Leptothorax. There is a single European species.

Revision: Smith, 1939. Ent. Soc. Wash., Proc. 41:165-172.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:26-28 (larvae).

Harpagoxenus americanus (Emery)
Mass., Ont. s. to N. C, w. to 111., Mo. Ecology: Enslaves Leptothorax
curvispinosus Mayr and L. longispinosus Roger.
Tomognathus americanus Emery, 1895. Zool. Jahrb., Abt. f. System 8: 272 3 .

Taxonomy: Creighton, 1927. Psyche 34:28 (male). —Smith, 1939. Ent. Soc. Wash., Proc.
41:166-168 (each caste). —Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:26-27
(larva).

Biology: Sturtevant, 1927. Psyche 34:1-9. —Creighton, 1927. Psyche 34:11-29. —Creighton,
1923. Psyche 36:48-50. —Wesson, 1939. Amer. Ent. Soc., Trans. 65:97-122.

Harpagoxenus canadensis Smith
N. S., Que., Maine w. to Mich., Minn. Ecology: Enslaves Leptothorax
muscorum (Nylander). Apparently closely related to the Palearctic H. sublaevis
(Nylander) and like that form has ergatoid females in addition to normal workers and
females.
Harpagoxenus canadensis Smith, 1939. Ent. Soc. Wash., Proc. 41:168. 9, ergatoid 9.

Taxonomy: Gregg, 1945. Canad. Ent. 77:74-76.




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Biology: Gregg, 1946. Amer. Midland Nat. 35:748.

Tribe MYRMECININI
Genus MYRMECINA Curtis

Myrmecina Curtis, 1829. Brit. Ent. 6:226, pi. 265.

Type-species: Myrmecina latreillii Curtis. Orig. desig.
Archaeomyrmex Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bui. 64:448, 451.

Type-species: Archaeomyrmex cacabau Mann. Orig. desig.

Apparently only one species occurs in North America, though several subspecies are some-
times recognized.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:271. —Smith, 1948. Ent. Soc. Wash.,
Proc. 50:238-240. — SnelUng, 1965. South. Calif. Acad. Sci., Bui. 64:101-105. —Brown, 1967.
Ent. News 78:233-230.

Taxonomy: Brown, 1951. Brooklyn Ent. Soc., Bui. 46:103-106. —Wheeler and Wheeler, 1954.
Ent. Soc. Wash., Proc. 56:129-131 (larvae). —Brown, 1971. Breviora 365:1-2 (generic syn.).

Myrmecina americana Emery
Que. s. to Ga., w. to Iowa, Colo., N. Mex., Ariz., Calif. Ecology: Colonies are

small and nests are obscure, usually built in moist shady areas often under small stones.
Myrmedtm latreillei americana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:271. ♂.
Myrmecina latreillei americana var. brevispinosa Emery, 1895. Zool. Jahrb., Abt. f.

System. 8:271. 5, 9,6.
Myrmecina graminicola texana Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:422. ♂.
Myrmeciria graminicola quadrispina Enzmann, 1946. N. Y. Ent. Soc., Jour. 54:13. ♂.
Myrmecina califomica Smith, 1948. Ent. Soc. Wash., Proc. 50:239. ♂.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24:39. — Buren, 1944. Iowa State Col., Jour. Sci.
18:290. —Brown, 1949. Psyche 56:44-47. —Brown, 1951. Brooklyn Ent. Soc., Bui.
46:103-106. —Wheeler and Wheeler, 1954. Ent. Soc. Wash., Proc. 56:130 (larva).
— Francoeur, 1966. Nat. Canad. 93:439.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:373, 376. —Wheeler, 1906. Amer. Mus.
Nat. Hist., Bui. 22:332, 335-336. —Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:502.
—Talbot, 1934. Ecology 15:420, 427^128. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:274,
278-279, 304. —Gregg, 1963. Ants of Colo., pp. 364-365, 367.

Tribe TETRAMORIINI

Revision: Bolton, 1976. Brit. Mus. (Nat. Hist.) Ent., Bui. 34: 283-379.

Genus TRIGLYPHOTHRIX Forel

Triglyphx)thrix Forel, 1890. Soc. Ent. de Belg., Ann. 34: CVI.
Type-species: Triglyphothrix walshi Forel. Monotypic.

A single introduced form is established in a number of localities in southeastern United
States. The genus is native to Africa and southern Asia.

Revision: Bingham, 1903 Fauna Brit. India, Hym. 2:172-175. —Bolton, 1976. Brit. Mus. (Nat.
Hist.) Ent., Bui. 34: 310-359 (world species).

Triglyphothrix lanuginosa (Mayr)
S. C, Ga., Fla., Ala, Miss., La.; pantropical. Introduced, probably native to
India. Spread by commerce to most tropical regions of the world.
Tetramorium lanuginosa Mayr, 1870. Zool.-Bot. GeseU. Wien, Verh. 20: 972, 976. ♂.
Tetramorium obesum race striatidens Emery, 1889. Ann. Mus. Civ. Stor. Nat. Genova
7:501. ♂.
Taxonomy: Wheeler, 1912. N. Y. Ent. Soc., Jour. 20:46. —Wheeler, 1916. Jour. Econ. Ent.
9:568-569 (first notice of occurrence in U. S.). —Wilson and Taylor, 1967. Pacific Ins.
Monog. 14:70. —Wheeler and Wheeler, 1973. Psyche 80: 78 Oarva). —Bolton, 1976. Brit.
Mus. (Nat. Hist.) Ent., Bui. 34: 350-352 (further synonymy).

Biology: Donisthorpe, 1927. British Ants, p. 393. —Smith, 1931. Ent. News 42:21.




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Genus TETRAMORIUM Mayr

Tetramorium Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:423.

Type-species: Formica caespitum Linnaeus. Desig. by Girard, 1879.
Tetrogmus Roger, 1857. Berlin. Ent. Ztschr. 1:10.

Type-species: Tetrogmus caldarius Roger. Monotypic.
Tetramorium subg. Xiphnmyrmex Forel, 1887. Schweiz. Ent. Gesell. Mitt. 7: 385.

Type-species: Tetramoriutn (Xiphomymiex) kelleri Forel. Desig. by Wheeler, 1911.

This genus is better represented in Asia and Africa than in North America. Three of the four
sp>ecies, guineense, pacificum,, and simillimum, are, with Uttle question, introduced; the other,
caespitum, which is also found in Europe, is generally believed to be native, though some wor-
kers also consider it as being introduced. This genus contains some important house pests.

Revision: Mayr, 1870. Zool.-Bot. GeseU. Wien, Verh. 20:972-977. —Bingham, 1903. Fauna
British India, Hym. 2:175-189. —Emery, 1916. Soc. Ent. Ital., Bol. 47:195. —Smith, 1938.
Wash. Acad. Sci., Jour. 28: 126-130 (Xiphomymiex). —Smith, 1943. Ent. Soc. Wash., Proc.
45:1-5 (U. S. species).

Taxonomy: Wheeler and Wheeler, 1954. Amer. Midland Nat. 52: 445, 450 (larvae). — Cole,
1957. Tenn. Acad. Sci., Jour. 32: 209-219 (male, Xiphomyrmex). —Brown, 1957. Breviora
72:1-8 (native in N. Amer. ?). —Bolton, 1976. Brit. Mus. (Nat. Hist.) Ent., Bui. 34: 288,
359-365 (generic synonymy).

Tetramorium caespitum (Linnaeus)
Ont. s. to Tenn., w. to Nebr., Mo., Wash., Nev., Calif.; Eurasia, Africa.

Ecology: One of the most common house-infesting ants in the large cities of the Atlantic
coast. They also steal seeds from seedbeds, gnaw into tubers, roots, and stalks of
various plants, attend honeydew excreting insects, and serve as an intermediate host for
poultry tapeworms. Colonies are populous and nests may be in exposed soil, under cover
of stones, pavement, or other objects, in rotting wood, or next to building foundations.
Possibly introduced by early colonists from Europe but believed by some to be a native
sp)ecies. Parasite: Anergates atratulus (Schenck), Strongylognathus sp. Pavement ant.
Most common in the Atlantic seaboard states and more sparsely distributed inland.
FoTvnica caespitum Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 581.

Myrmica (Myrmica) brevinodis var. transversinodis Enzmann, 1946. N. Y. Ent. Soc., Jour.
54:47. ♂.

Taxonomy: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:972. — Donisthorpe, 1927. British
Ants, p. 189 (each caste). —Smith, 1943. Ent. Soc. Wash., Proc. 45:2. —Brown, 1949. Psyche
56:47. — Wheeler and Wheeler, 1954. Amer. Midland Nat. 52:445 (larva). — Wheeler and
Wheeler, 1960. Ent. Soc. Wash., Proc. 62:24-25 Oarva). —Wheeler and Wheeler, 1973.
Psyche 80: 76, 78 (larva).

Biology: Smith, 1915. Va. Truck Expt. Sta., Bui. 16:1-15. —Wheeler, 1919. Amer. Phil. Soc.,
Proc. 58:23-26. —Wheeler, 1927. Psyche 34:164-165. —Donisthorpe, 1927. British Ants, pp.
193-198. — Metcalf and Flint, 1939. Destructive and Useful Insects, p. 771. — Mallis, 1941.
South. Calif. Acad. Sci., Bui. 40:74. —Brown, 1964. Ent. News 75:15. —Brown, 1957.
Breviora 72:1-4 (historical). —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:45-47
(economic importance). — Lange, 1961. Jour. Econ. Ent. 54:1063. — Poldi, 1963. Pavia Univ.
Symposium Genet, et Biol. Ital. 12:132. —Weber, 1965. Ent. News 76: 137-139 (in
Philadelphia area). —Brian, Elmes, and Kelley, 1967. Jour. Anim. Ecol. 36:337-342
(populations). — Bruder and Gupta, 1972. Ent. Soc. Amer., Ann. 65:358-367. — Gumey, 1975.
Ins. World Digest 2 (5): 19-26 (stinging habits).

Tetramorium guineense (Fabricius)
Ga., Fla. w. to Tex.; pantropical. Ecology: Nests in smaU to moderate

colonies in exposed soil, under stones or other objects, in rotting logs and stumps, in

stems of plants, and in branches and under bark of trees. Occasionally a house pest.

Introduced, spread throughout the tropical regions of the world by commerce.

Occasionally found in greenhouses farther north in N. Amer. Guinea ant.
Formica guineensis Fabricius, 1793. Ent. System. 2:357. ♂.

Taxonomy: Emery, 1909. Deut. Ent. Ztschr., p. 695 (each caste). —Smith, 1936. Puerto Rico
Univ., Jour. Agr. 20:831, 852. —Smith, 1943. Ent. Soc. Wash., Proc. 45:3. —Wilson and




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Taylor, 1967. Pacific Ins. Monog. 14:71-72 (Polynesia). —Wheeler and Wheeler, 1954. Amer.
Midland Nat. 52:449-450 (larva).

Biology: Marlatt, 1928. U. S. Dept. Agr., Farmers' Bui. 740:6. —Phillips, 1934. (Hawaii Univ.)
Expt. Sta. Pineapple Prod. Coop. Assoc. Bui. 15:23-24. —Smith, 1943. Ent. Soc. Wash.,
Proc. 45:1-2. —Brown, 1958. Acta Hym. 1:28. —Taylor and Wilson, 1961. Psyche 68:138.
—Brown, 1964. Ent. News 75:14-15. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:47-48
(economic importance).

Morphology: Blum and Ross, 1965. Jour. Ins. Physiol. 11:857-868 (odor trail pheromone).

Tetramorium pacificum Mayr
Calif.; Oriental and Australian Regions. Introduced into a nursery in Calif;
may not be established.
Tetramarium pacificum Mayr, 1870. Zool.-Bot. (Jesell. Wien, Verb. 20:972, 976. 9,9.

Taxonomy: Smith, 1943. Ent. Soc. Wash., Proc. 45:2-3. —Wilson and Taylor, 1967. Pacific Ins.
Monog. 14:72-73 (Polynesia).

Tetramorium simillimum (Smith)
Ga., Fla.; Pantropical. Introduced, a tramp species probably native to
Africa. Occasionally found in greenhouses farther north in N. Amer.
Myrmica simillima Smith, 1851. List Hym. Brit. Mus. v. 6, p. 118. ♂.

Taxonomy: Emery, 1909. Deut. Ent. Ztschr., pp. 695-696 (each caste). —Smith, 1936. Puerto
Rico Univ., Jour. Agr. 20:831, 853. —Smith, 1943. Ent. Soc. Wash., Proc. 45:2. —Wilson and
Taylor, 1967. Pacific Ins. Monog. 14:73 (Polynesia).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:101. —Marlatt, 1928. U. S. Dept. Agr.,
Farmers' Bui. 740:6. —Smith, 1933. Fla. Ent. 17:24. —Phillips, 1934. (Hawaii Univ.) Expt.
Sta. Pineapple Prod. Coop. Assoc. Bui. 15:24. —Taylor and Wilson, 1961. Psyche 68:142-143.
spinosus hispidus (Wheeler). S. Ariz. Ecology: Found nesting in small craters in the desert. X.
spinosus spinosus Pergande occurs in Mexico.
Xiphomyrmex spinosus hispidus Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:415. ♂.
spinosus insons (Wheeler). W. Tex., Ariz. Ecology: Nests found in small craters in dry, grassy
areas.
Xiphomyrmex spinosus insons Wheeler, li915. Amer. Mus. Nat. Hist., Bui. 34:416. 9, 9, d.
spinosus wheeleri Forel. S. Ariz.; Mexico. Ecology: One colony was found beneath a stone in a
cactus desert.
Tetramorium (Xiphomyrmex) Wheeleri Forel, 1901. Soc. Ent. de Belg., Ann. 45:128. ♀.

Biology: Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:416.

Genus ANERGATES Forel

Anergates Forel, 1874. Schweiz. Naturf. Gesell. Denkschr. 26:93.
Type-species: Myrmica atratula Schenck. Monotypic.

The single species of this genus is a permanent parasite.

Anergates atratulus (Schenck)
Conn., N. Y., N. J., Pa., Del., Md., D. C, Va.; Europe. Ecology: A

parasitized nest of the host consists of a single fertile female of atratulus, a large
number of host workers, and a large number of pupoidal males and virgin females of
atratulus. Workers of the parasite are entirely lacking. Possibly introduced with its host
from Europe, though some workers are of the opinion that the host and atratulus are
native. Host: Tetramorium caespitum, (L.).

Myrmica atratula Schenck, 1852. Nassau. Ver. f. Naturk. Jahrb. 8:91. ♀.

Anergates friedlandi Creighton, 1934. Psyche 41:193. ♀.

Taxonomy: Emery, 1922. In Wytsman, Gen. Ins., fasc. 174:205-206 (female, male).

— Donisthorpe, 1927. British Ants, pp. 96-97 (female, male). —Wheeler and Wheeler, 1955.
Amer. Midland Nat. 54:128-130 (larva). — Ettershank, 1966. Austral Jour. Zool. 14:157-158.

Biology: Wheeler, 1910. Ants, pp. 498-504. —Donisthorpe, 1915. British Ants, p. 89.

—Wheeler, 1923. Social life Among the Insects, pp. 215-219. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bui. 104:241-243.

Morphology: Meyer, 1955. Insectes Sociaux 2:163-170.




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Tribe OCHETOMYRMICINI
Genus OCHETOMYRMEX Mayr

Ochetomyrmex Mayr, 1877. Zool.-Bot. Gesell. Wien, Verb. 27:871.

Type-species: Ochetomyrmex semipolitus Mayr. Monotypic.
Wasmannia Forel, 1893. London Ent. Soc, Trans. 4:383.

Type-species: TetramoriumY auropunctatum Roger. Desig. by Wbeeler, 1911.
Blepharidatta Wbeeler, 1915. Harvard Univ., Mus. Comp. ZooL, Bui. 59:484.

Type-species: Blepharidatta brasiliensis Wbeeler. Monotypic.
Hercynia Enzmann, 1947. N. Y. Ent. Soc, Jour. 55:43. Preocc. by Rafinesque, 1815.

Type-species: Hercynia panamana Enzmann. Monotypic.

A single species of tbis Neotropical genus bas been introduced into the United States.

Taxonomy: Brown, 1948. Ent. News 59:102. —Brown, 1973. In Meggers, et al.. Tropical
Forest Ecosystems in Afr. and S. Amer., pp. 178-185 (generic syn.).

Ochetomyrmex auropunctata (Roger)
Fla., Calif.; W. Indies, Mexico, Central and S. Amer. Ecology: Nests in
exposed soil, under cover of objects, or in wood. Attends boneydew excreting insects,
can sting severely, and infests houses. Introduced. Little fire ant.
Tetramorium ? auropunctatum Roger, 1863. Berlin. Ent. Ztschr. 7:182. 5, 9, 6.
Wasmnnnia ^ glabra Santscbi, 1931. Rev. Ent. Sao Paulo 1:272. ♂.
Hercynia panamana Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:44. 5,2.

Taxonomy: Smith, 1929. Jour. Econ. Ent. 22:243. —Smith, 1936. Puerto Rico Univ., Jour. Agr.
20:831, 854. —Brown, 1948. Ent. News 59:102. —Smith, 1954. Amer. Mus. Novitates
1671:7-8. —Wbeeler and Wheeler, 1954. Amer. Midland Nat. 52:444 (larva). — Kempf, 1964.
Studia Ent. 7:66.

Biology: Wbeeler, 1929. Psyche 36:89-90. —Spencer, 1941. Fla. Ent. 24:6-14. — Femald, 1947.
Jour. Econ. Ent. 40:428. — Osbum, 1948. Fla. Ent. 31:11-15. —Smith, 1965. U. S. Dept.
Agr., Tech. Bui. 1326:48-49 (economic importance).

Tribe CEPHALOTINI

Revision: Kempf, 1951. Rev. de Ent. 22:1-244.

Taxonomy: Kempf, 1958. Studia Ent. 1:1-168. —Kempf, 1973. Studia Ent. 16: 449-462 (generic
classification; key to genera).

Genus ZACRYPTOCERUS Wheeler

Zacryptocenis Asbmead, 1905. Canad. Ent. 37: 384. Nomen nudum. No description given;
the type-sp»ecies is given as Cryptocerus multistrigus F. Smith, but no such species
exists.
Zacryptocenis Wheeler, 1911. N. Y. Acad. Sci., Ann. 21: 175.

Type-species: Cryptocerus clypeatus Fabricius. Monotypic.
Cryptocerus subg. Paracryptocerus Emery, 1915. Soc. Ent. de France, Bui., p. 192.

Type-species: Cryptocerus spinosus Mayr. Orig. desig.
Cryptocerus subg. Cyathocephalus Emery, 1915. Soc. Ent. de France, Bui., p. 192. Preocc.
by Kessler, 1868.

Type-species: Cryptocerus pallens Klug. Orig. desig.
Cryptocerus subg. Hypocryptocerus Wheeler, 1920. Psyche 27: 53.

Type-species: Formica haemorrhoidalis Latreille, Orig. desig.
Cyathomyrmex Creighton, 1933. Psyche 40:98. N. name for Cyathocephalus Emery.
Paracryptocerus subg. Hamedia Smith, 1949. Psyche 56:20.

Type-species: Cryptocerus umbraculatus Fabricius. Orig. desig.

Mainly Neotropical with slight extensions into the United States. The ants are arboreal, form-
ing small colonies in cavities of plants, especially twigs. Food consists largely of boneydew and
small arthropods. The soldier is believed to keep intruders from the nest by blocking the en-
trance hole with its bead.

Revision: Smith, 1947. Ent. Soc. Wash., Proc. 49:29-40. —Kempf, 1958. Studia Ent. 1:65-158.




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Taxonomy: Kempf, 1951. Rev. de Ent. 22:156-157, 233. —Wheeler and Wheeler, 1954. Wash.
Acad. Sci., Jour. 44:156-157 (larvae). —Kempf, 1973. Studia Ent. 16: 457-460 (syn. of
Paracryptocerus and Hypocryptocerus).

Zacryptocerus rohweri (Wheeler)
n. comb. S. Ariz.; Mexico. Ecology: Arboreal.

Cryptocerus (Cyathocephalus) rohweri Wheeler, 1916. New England Zool. Club, Proc. 6:32.

Taxonomy: Kempf, 1958. Studia Ent. 1:129-132. — Snelling, 1968. Los Angeles Co., Mus.,
Contrib. Sci. 132:5-9.

Biology: Smith, 1947. Ent. Soc. Wash., Proc. 49:34-37. — Creighton and Nutting, 1965. Psyche
72: 59-64 (habits and distribution).

Zacryptocerus texanus (Santschi)
n. comb. S. Tex.; Mexico. Ecology: Arboreal.

Cryptocerus texanus Santschi, 1915. Soc. Ent. de France, Bui., p. 208. 9, %.

Taxonomy: Kempf, 1958. Studia Ent. 1:123-127.

Biology: Smith, 1947. Ent. Soc. Wash., Proc. 49:37-40. —Creighton, 1954. Psyche 61:41-57.
—Creighton, 1963. Psyche 70:133-143.

Zacryptocerus varians (Smith)
n. comb. Fla.; Central Amer., W. Indies. Ecology: Arboreal.
Cryptocerus varians Smith, 1876. London Ent. Soc., Trans., p. 606. ♂.
Cryptocerus (Cyathocephalus) varians var. jamaicensis Forel, 1922. Rev. Suisse de Zool.
30:97. ♂.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:102-104. —Wheeler and Wheeler,
1954. Wash. Acad. Sci., Jour. 44:156 (larva). —Kempf, 1958. Studia Ent. 1:155-158.

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:11. —Smith, 1947. Ent. Soc. Wash., Proc.
49:33.

Tribe DACETINI

Taxonomy: Brown, 1948. Amer. Ent. Soc., Trans. 74:101-129 (preUminary generic revision of
higher Dacetini). —Brown, 1949. Amer. Ent. Soc., Trans. 75:43-51. — Brovm, 1953. Amer.
Midland Nat. 50:1-137 (revisionary studies). —Wheeler and Wheeler, 1954. Psyche
61:111-145 Garvae). —Brown and Wilson, 1959. Quart. Rev. Biol. 34:278-294 (evolution of
dacetine ants).

Genus STRUMIGENYS Smith

Strumigenys Smith, I860. Jour. Ent. (London) 1:72.

Type-species: Strumigenys mandibularis Smith. Monotypic.
Labidogenys Roger, 1862. Berlin. Ent. Ztschr. 6:249.

Type-species: Labidogenys lyroessa Roger. Monotypic.
Pyramica Roger, 1862. Berlin. Ent. Ztschr. 6:251.

Type-species: Pyramica gundlachi Roger. Monotypic.
Proscopomyrmex Patrizi, 1946. Bol. Inst. Ent. R. Univ. Bologna 15:294.

Type-species: Proscopomyrmex londianensis Patrizi. Monotypic.
Eneria Donisthorpe, 1947. Ann. and Mag. Nat. Hist. 14:598.

Type-species: Eneria excisa Donisthorpe. Monotypic.

This genus is best represented in the tropical regions of the world; over 50 sjjecies are known
from the New World, but only four of these have been recorded from the United States. Three
of these four species are believed to have been introduced. Colonies are small and nests are
usuaUy found in leaf litter, plant cavities, rotting wood, or under objects.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325-326. —Wheeler, 1908. Amer. Mus.
Nat. Hist, Bui. 24:145-147. —Smith, 1931. Ent. Soc. Amer., Ann. 24:688^91 (N. Amer.).
— Brovm, 1962. Psyche 69:238-267 (Neotropical species and key).

Taxonomy: Brown, 1948. Amer. Ent. Soc., Trans. 74:108. —Brown, 1949. Mushi 20:14.

—Wheeler and Wheeler, 1954. Psyche 61:135 Oarvae). —Wheeler and Wheeler, 1960. Ent.
Soc. Wash., Proc. 62:25-26 (larvae). —Brown, 1959. Studia Ent. (n. s.) 2:25-30 (silvestrii
group). —Brown, 1960 (1959). Psyche 66:37-52 (gundlachi group).




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Biology: Brown, 1950. Brooklyn Ent. Soc, Bui. 45:87-89. —Brown and Wilson, 1960 (1959).
Quart. Rev. Biol. 34:281.

Species Group Silvestrii

Strumigenys silvestrii Emery
La. (Paradise); Brazil, Argentina, Cuba. Ecology: In La. found in a basal rot
hole in a live oak tree. Probably introduced.
Strumigenys silvestrii Emery, 1905. Soc. Ent. Ital., Bol. 37:1()8. ♂, ♀.
Strumigenys {Strumigenys) caribbea Weber, 1934. Rev. de Ent. 4:43. ♂, ♀.

Taxonomy: Brown, 1959. Studia Ent. (n. s.) 2:25-28. —Brown, 1962. Psyche 69:246, 257, 259.

Species Group Louisianae

Strumigenys louisianae Roger
N. C. to Fla., w. to Okla., Ariz.; Mexico s. to Bolivia, Argentina. Ecology:

The small colonies are beneath objects, in rotting wood, plant cavities; food consists of

small arthropods, especially Collembola.
Strumigenys louisianae Roger, 1863. Berlin. Ent. Ztschr. 7:211. ♂.
Strumigenys unidentata Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:570, 575. ♂.
Strumigenys unispinulosa Emery, 1890. Soc. Ent. Ital., Bol. 22:67. ♂, ♀.
Strumigenys unispinulosa var. longicomis Emery, 1894. Soc. Ent. Ital., Bol. 26:214. ♂.
Strumigenys fusca Emery, 1894. Soc. Ent. Ital., Bol. 26:215. ♂.
Strumigenys louisianae var. obscuriventris Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.

24:145. ♂.
Strumigenys bruchi Forel, 1912. Soc. Ent. de Belg., Mem. 29:197-198. ♀, ♂.
Strumigenys injidelis Santschi, 1919. Soc. Cient. Argentina, An. 87:48. ♂.
Strumigenys eggersi var. cubaensis Mann, 1920. Amer. Mus. Nat. Hist., Bui. 42:430. ♂.
Strumigenys (Strumigenys) louisianae laticephala Smith, 1931. Ent. Soc. Amer., Ann.

24:688, 690. ♂.
Strumigenys (Strumigenys) louisianae soledadensis Weber, 1934. Rev. de Ent. 4:38-39. ♂.

9.
Strumigenys (Strumigenys) louisianae guatemalensis Weber, 1934. Rev. de Ent. 4:39. ♂.
Strumigenys (Strumigenys) louisianae costaricensis Weber, 1934. Rev. de Ent. 4:39. ♂.
Strumigenys clasmospongia Brown, 1953. Psyche 60:2. § .

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325-326. —Wheeler, 1908. Amer.
Mus. Nat. Hist., Bui. 24:147. — Haug, 1932. Ent. Soc. Amer., Ann. 25:170-172. —Brown,

1953. Amer. Midland Nat. 50:28-31. —Brown, 1953. Psyche 60:1-2. —Wheeler and Wheeler,

1954. Psyche 61:136-137 (larva). —Brown, 1957. Quart. Rev. Biol. 32:271. —Brown, 1961.
Psyche 68: 64-67, 68 (variation and syn.). —Brown, 1962. Psyche 69:246-247, 257, 263.

Biology: Smith, 1931. Ent. Soc. Amer., Ann. 24:690-691. — Creighton, 1937. Psyche 44:97-109.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31:269, 291, 305. — WUson, 1950. Brooklyn Ent. Soc.,
Bui. 45:85-86. —Wilson, 1953. Ent. Soc. Amer., Ann. 46:481-483. — Kempf, 1958. Studia Ent.
1:555.

Species Group Gundlachi

Strumigenys eggersi Emery
Fla. (Miami, Fisher's Is., Archbold Biol. Sta.); W. Indies, Mexico, S. Amer.
Probably introduced.
Strumigenys eggersi Emery, 1890. Soc. Ent. Ital., Bol. 22:68. ♂, ♀.

Taxonomy: Brown, 1960 (1959). Psyche 66:46-47 (worker, female). —Brown, 1962. Psyche
69:249, 257, 264.

Biology: Weber, 1952. Amer. Mus. Novitates 1554:5. —Kempf, 1958. Studia Ent. 1:554.
— Kempf, 1961. Studia Ent. 4:515.

Strumigenys gundlachi (Roger)
Fla. (Royal Palm Ranger Sta., Everglades Natl. Pk.; Key Largo); W.
Indies, Mexico, Central Amer. Ecology: Usually found in leaf litter; feeds almost
exclusively on Collembola. Probably introduced.
Pyramica gmidlachi Roger, 1862. Berlin. Ent. Ztschr. 6:253. "♀"=♂.
Strumigenys eggersi var. vincentensis Forel, 1893. London Ent. Soc., Trans, p. 378. ♂.




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Strumigenys eggersi var. banillensis Santschi, 1930. Soc. Roy. Ent. d'Egypte, Bui. (n. s.), p.

80. ♂.
Strumigenys bierigi Santschi, 1930. Soc. Roy. Ent. d'Egypte, Bui. (n. s.), p. 80. ♂.
Strumigenys (Strumigenys) eggersi var. isthmica Santschi, 1931. Rev. de Ent. 1:276. ♂.
Strumigenys (Strumigenys) eggersi infuscata Weber, 1934. Rev. de Ent. 4:35. ♂, ♀.
Strumigenys (Strumigenys) eggersi var. berlesei Weber, 1934. Rev. de Ent. 4:36. ♀.

Taxonomy: Brown, 1960 (1959). Psyche 66:37-45 (worker, female). —Brown, 1962. Psyche
69:249, 257, 264.

Biology: Weber, 1952. Amer. Mus. Novitates 1554:4-5.

Genus SMITHISTRUMA Brown

Cephaloxys Smith, 1864. Linn. Soc. London, Jour. 8:77. Preocc. by Signoret, 1847.

Type-species: Cephaloxys capitata Smith. Monotypic.
Smithistruma Browm, 1948. Amer. Ent. Soc., Trans. 74:104-106.

Type-species: Strum-igenys pulchella Emery. Orig. desig.
Smithistruma subg. Wessonistruma Brovm, 1948. Amer. Ent. Soc., Trans. 74:106.

Type-species: Strumigenys pergandei Emery. Orig. desig.
Smithistruma subg. Weberistruma Brown, 1948. Amer. Ent. Soc., Trans. 74:106-107.

Type-species: Strumigenys (Cephaloxys) leptothrix Wheeler. Orig. desig.
Smithistruma subg. Platystruma Browrn, 1953. Amer. Midland Nat. 50:112.

Type-species: Strumigenys (Cephaloxys) depressiceps Wheeler. Orig. desig.

Ants of this genus nest in small colonies in the soil, various tyi)es of soil cover, rotting logs and
stumps, and beneath objects. They are most frequently collected in berlese funnels of samples of
leaf litter, duff, etc. Food consists almost exclusively of Collembola.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:32&-329. —Wheeler, 1908. Amer. Mus.
Nat. Hist, Bui. 24:148. —Smith, 1931. Ent. Soc. Amer., Ann. 24:691-710. —Wesson and
Wesson, 1939. Psyche 46:91-112. —Brown, 1953. Amer. Midland Nat. 50:55-92 (key to spp.).

Taxonomy: Wheeler and Wheeler, 1954. Psyche 61:141-145 (larvae).

Biology: Brown, 1950. Brooklyn Ent. Soc., Bui. 45:87-89. —Wilson, 1953. Ent. Soc. Amer.,
Ann. 46:485-487. —Brown and Wilson, 1960 (1959). Quart. Rev. BioL 34:282. —Brown, 1964.
Amer. Ent. Soc., Trans. 89:183-200.

Smithistruma abdita (Wesson and Wesson)
Pa., Va., N. C, Ohio, 111., Iowa. Ecology: Nests are found under
rocks in the soil.
Strumigenys (Cephaloxys) abdita Wesson and Wesson, 1939. Psyche 46:95, 109. ♀.

Taxonomy: BrowTi, 1953. Amer. Midland Nat. 50:89-90 (worker, female). —Brown, 1964.
Amer. Ent. Soc., Trans. 89:189-190, 199-200.
angfulata (Smith). 111., Ala., Miss. Ecology: Colonies fovmd in a rotten pine stump; some
specimens found in nest of Strumigenys louisianae Roger.
Strumigenys (Cephaloxys) angulata Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 697. ♂.

Taxonomy: Smith, 1932. Ent. News 43:157-158 (also biology). —Wesson and Wesson, 1939.
Psyche 46:110. — Brown, 1953. Amer. Midland Nat. 50:54-55 (worker, female).
bimarg^nata (Wesson and Wesson). Ohio, 111. Ecology: Specimens found in soil cover.

Strumigenys (Cephaloxys) bimarginata Wesson and Wesson, 1939. Psyche 46:95-97. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:62-63 (worker, female).

Smithistruma bunki Brown
Ga., Fla, Miss., La. Ecology: Nests are found in the soiL

Smithistruma (Smithistruma) bunki Brown, 1950. Amer. Ent. Soc., Trans. 76:41-42. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:82-84 (worker, female). — Brown, 1964.
Amer. Ent. Soc., Trans. 89:190.
califomica Brown. Calif.

Smithistruma (Smithistruma) califomica Brown, 1950. Amer. Ent. Soc., Trans. 76:40-41.

9.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:85-86.




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Smithistruma carolinensis Brown
N. C, S. C. Ecology: Specimens found in pine needle litter and oak leaf
mold.

Smithistruma carolinensis Brown, 1964. Amer. Ent. Soc, Trans. 89:185-186. ♂.

Smithistruma cloydi Pfitzer
Tenn. (Knoxville).

Smithistruma cloydi Pfitzer, 1951. Tenn. Acad. Sci., Jovtr. 16:198-200. ♂.

Taxonomy: Brown, 1964. Amer. Ent. Soc, Trans. 89:191.

Smithistruma clypeata (Roger)
N. J., Pa. s. to Fla., w. to 111., Ark., La. Ecology: Usually found in forested
areas where it nests in soil cover, under stones, and in rotten logs.
Strumigenys clipeata(\) Roger, 1863. Berlin. Ent. Ztschr. 7:213. ♂.

Taxonomy: Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:571 (worker, female). — Emery,
1895. Zool. Jahrb., Abt. f. System. 8:326, 328 (each caste). —Wheeler, 1908. Amer. Mus.
Nat. Hist., Bui. 24:148. —Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 699-700. —Brown,

1953. Amer. Midland Nat. 50:58-59 (each caste). — Wheeler and Wheeler, 1954. Psyche
61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:191 (further distribution).

Biology: Smith, 1932. Ent. News 43:159. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:291, 305.
—Wesson and Wesson, 1939. Psyche 46:93-94. —Smith, 1955. Brooklyn Ent. Soc., Bui.
50:28. —Wilson, 1953. Ent. Soc. Amer., Ann. 46:486.

Smithistruma creightoni (Smith)
D. C, N. C, Ga., Fla., Tenn., Ala. Ecology: Collections made from leaf
Utter.
Strum.igenys (Cephaloxys) creightoni Smith, 1931. Ent. Soc. Amer., Ann. 24:692, 705. ♂.

Taxonomy: Wesson and Wesson, 1939. Psyche 46:110. — Brown, 1953. Amer. Midland Nat.
50:80-81 (worker, female). —Brown, 1964. Amer. Ent. Soc., Trans. 89:191.

Smithistruma dietrichi (Smith)
Md. s. to Fla., w. to Ohio, 111., Ark., La. Ecology: Nests in or imder bark of
rotting logs or stumps; a few are known from leaf litter. Sometimes occurs in or near
nests of other ants.
Strumigenys {Cephaloxys) dietrichi Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 696. ♀.

Taxonomy: Smith, 1932. Ent. News 43:159. —Wesson and Wesson, 1939. Psyche 46:108.
— Brown, 1953. Amer. Midland Nat. 50:67-68 (worker, female). — Wheeler and Wheeler,

1954. Psyche 61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:191-192.

Biology: Kennedy and Schramm, 1933. Ent. Soc. Amer., Ann. 26:95-96. —Dennis, 1938. Ent.
Soc. Amer., Ann. 31:270, 272-273, 291, 305. —Wilson, 1953. Ent. Soc. Amer., Ann. 46:487.

Smithistruma filirrhina Brown
N. C, Mo. Ecology: Specimens taken in leaf mold.

Smithistruma {Smithistruma) filirrhina Brown, 1950. Amer. Ent. Soc., Trans. 76:37-38. 2 .

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:63-64. —Brown, 1964. Amer. Ent. Soc.,
Trans. 89:193.

Smithistruma filitalpa Brown
Ind., Ark.

Smithistruma {Smithistruma) filitalpa Brown, 1950. Amer. Ent. Soc., Trans. 76:39-40. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:79-80. —Brown, 1964. Amer. Ent. Soc.,
Trans. 89:193-194.

Smithistruma laevinasis (Smith)
Va., N. C, Tenn., Ala., Miss., 111. Ecology: Nests have been found in rotten
logs.
Strumigenys {Cephaloxys) clypeata var. Uievinasis Smith, 1931. Ent. Soc. Amer., Ann.
24:691, 701. ♀.

Taxonomy: Wesson and Wesson, 1939. Psyche 46:109. — Brown, 1953. Amer. Midland Nat.
50:61-62 (worker, female). —Brown, 1964. Amer. Ent. Soc., Trans. 89:194, 199.

Biology: Smith, 1932. Ent. News 43:159.

Smithistruma margaritae (Forel)
Ga., Ala., Tex.; W. Indies, Mexico, Colombia Ecology: Specimens have
been found in soil cover.
Strumigenys margaritae Forel, 1893. London Ent. Soc., Trans, p. 378. 5, 9, 6.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:148. —Smith, 1931. Ent. Soc.
Amer., Ann. 24:691, 692-693. —Wesson and Wesson, 1939. Psyche 46:108. —Brown, 1953.
Amer. Midland Nat. 50:108-110. —Brown, 1964. Amer. Ent. Soc., Trans. 89:194-195.




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Smithistruma missouriensis (Smith)
N. Y., Va., N. C, Ohio, Miss., 111., Iowa, Mo. Ecology: Primarily a soil
and soil cover inhabitant.
Strumigenys {Cephaloxys) missouriensis Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 701.

5.
Strumigenys {Cephaloxys) sculpturata Smith, 1931. Ent. Soc. Amer., Ann. 24:692, 706. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:72-73 (worker, female). —Wheeler and
Wheeler, 1954. Psyche 61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:195.

Biology: Wilson, 1953. Ent. Soc. Amer., Ann. 46:485-486.

Smithistruma ohioensis (Kennedy and Schramm)
N. J. s. to Ga., w. to III, Ark., La Ecology: Found in soil
cover and upper soil layers, often under objects; sometimes in rotting wood.
Strumigenys ohioensis Kennedy and Schramm, 1933. Ent. Soc. Amer., Ann. 26:98-99. ♂.
Strumigenys (Cephaloxys) manni Wesson and Wesson, 1939. Psyche 46:97. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:87-89 (each caste). —Brown, 1964. Amer.
Ent. Soc, Trans. 89:196.

omata (Mayr). Del, Md. s. to Fla., w. to Mich., Ill, Mo., Okla., Tex. Ecology: Most commonly
found in leaf litter or forest debris.
Strumigenys omata Mayr, 1887. Zool.-Bot. Gesell. Wien, Verb. 37:571. ♂.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325, 328. —Wheeler, 1908. Amer.
Mus. Nat. Hist., Bui. 24:148 (worker, female). —Smith, 1931. Ent. Soc. Amer., Ann. 24:691,
695-696. —Wesson and Wesson, 1939. Psyche 46:109. —Brown, 1953. Amer. Midland Nat.
50:65-67. —Brown, 1964. Amer. Ent. Soc., Trans. 89:197.

Smithistruma pergandei (Emery)
Mass., N. Y., Ont. s. to N. C, Tenn., w. to Iowa, Mo., Kans. Ecology: Nests
in rotten logs, in soil, or under stones in soil. Often found in or near nests of other ants
where it may prey on myrmecophilous collembolans. Records from Mass. are the
northernmost for dacetine ants in the New World.
Strumigenys pergandei Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325. 5, 9, i.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:148. —Smith, 1931. Ent. Soc.
Amer., Ann. 24:691, 698-699 (also biology). —Wesson and Wesson, 1939. Psyche 46:110.
— Brown, 1953. Amer. Midland Nat. 50:51-54. — Wheeler and Wheeler, 1954. Psyche
61:144-145 (larva). —Brown, 1964. Amer. Ent. Soc, Trans. 89:197.

Biology: Kennedy and Schramm, 1933. Ent. Soc. Amer., Ann. 26:95-98. —Wesson, 1936. Ent.
News 47:171-174. —Brown, 1952. Psyche 59:12. —Wilson, 1953. Ent. Soc. Amer., Ann.
46:487^88.

Smithistruma pilinasis (Forel)
Pa. s. to N. C, Ala., w. to Ohio., 111., Mo., Ark., La. Ecology: Nests imder
stones, in soil cover, or in logs.
Strumigenys clypeata var. pilinasis Forel, 1901. Soc. Ent. de Belg., Ann. 45:339. 2 .
Strumigenys (Cephaloxys) clypeata var. brevisetosa Smith, 1935. Ent. Soc. Amer., Ann.

28:215. ♀.
Strumigenys (Cephaloxys) medialis Wesson and Wesson, 1939. Psyche 46:94, 110. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:148 (worker, female). —Smith,
1931. Ent. Soc. Amer., Ann. 24:691, 700. —Wesson and Wesson, 1939. Psyche 46:109.
—Brown, 1953. Amer. Midland Nat. 50:60-61 (worker, female). —Wheeler and Wheeler,
1954. Psyche 61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:197-199.

Biology: Wilson, 1953. Ent. Soc. Amer., Ann. 46:485.

Smithistruma pulchella (Emery)
N. Y., Pa. s. to Fla., w. to 111., La. Ecology: Nests in rotting wood.
Strumigenys pulchella Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325, 327. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:148 (worker, female). —Smith,
1931. Ent. Soc. Amer., Ann. 24:691, 702. —Brown, 1953. Amer. Midland Nat. 50:70-72 (each
caste). —Brown, 1964. Amer. Ent. Soc., Trans. 89:198.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:291, 305. —Wesson and Wesson, 1939.
Psyche 46:111.




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Smithistruma reflexa (Wesson and Wesson)
Md. s. to N. C, Tenn., Ala., w. to Ohio, 111. Ecology: Nests in soil
or under or in objects lying on soil.
Strumigenys (Cephaloxys) reflexa Wesson and Wesson, 1939. Psyche 46:102, 111. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:73-75 (each caste).

Smithistruma rohweri (Smith)
Miss.

Strumigenys (Cephaloxys) rohweri Smith, 1935. Ent. Soc. Amer., Ann. 28:214. ♂.

Taxonomy: Wesson and Wesson, 1939. Psyche 46:109. — Brown, 1953. Amer. Midland Nat.
50:57-58.

Smithistruma rostrata (Emery)
N. J., Pa. s. to Fla., w. to 111., Mo., La. Ecology: Nests in soil and rotting
wood.
Strumigenys rostrata Emery, 1895. Zool. Jahrb., Abt. f. System. 8:326, 329. 5, 9, S.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:148. —Smith, 1931. Ent. Soc.
Amer., Ann. 24:692, 704-705. —Wesson and Wesson, 1939. Psyche 46:110. —Brown, 1953.
Amer. Midland Nat. 50:84-85 (each caste). — Wheeler and Wheeler, 1954. Psyche 61:144
(larva).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:273, 291-292, 305. —Wilson, 1953. Ent. Soc.
Amer., Ann. 46:485.

Smithistruma talpa (Weber)
N. C, Fla., Ohio, Tenn., Ala., 111., La. Ecology: Found in soil or humus.
Strumigenys (Cephaloxys) talpa Weber, 1934. Psyche 41:63. ♂.
Strumigenys (Cephaloxys) venatrix Wesson and Wesson, 1939. Psyche 46:103, 110. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:76-78 (each caste). —Wheeler and Wheeler,
1954. Psyche 61:141 (larva).

Biology: Wilson, 1953. Ent. Soc. Amer., Ann. 46:486.

Smithistruma wrayi Brown
N. C. (Fayetteville). Ecology: Found in leaf mold.

Smithistruma (Smithistruma) vrrayi Brown, 1950. Amer. Ent. Soc., Trans. 76:38-39. ♂.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:78-79.

Genus TRICHOSCAPA Emery

Strumigenys subg. Trichoscapa Emery, 1869. Accad. Degli Aspiranti Napoli, Ann. 2:24.
Type-species: Strumigenys (Trichoscapa) membranifera Emery. Monotypic.

The single species in this genus has been spread throughout the tropical regions of the world
by commerce.

Revision: Brown, 1948. Amer. Ent. Soc., Trans. 74:112-114.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:144-145, 148. —Smith, 1931. Ent.
Soc. Amer., Ami. 24:691, 693-695.

Biology: Brown and Wilson, 1960 (1959). Quart. Rev. Biol. 34:283.

Trichoscapa membranifera (Emery)
Ga., Fla., Ala., Miss., La., Calif, (nr. Stockton); Pantropical. Ecology:

Nests are in soil, wood, and plant cavities, and the ants are predaceous on a variety of

small soft-bodied arthropKxls. Introduced, probably of African origin.
Strumigenys (Trichoscapa) mem,branifera Emery, 1869. Accad. Degli Aspiranti Napoli,

Ann. 2:24. ♂.
Strumigenys membranifera sim.illima Emery, 1890. Soc. Ent. Ital., Bol. 22:69. ♂.
Strumigenys membranifera var. santschii Forel, 1904. Rev. Suisse de Zool. 12:6. ♂.
Strumigenys (Cephaloxys) vitiensis Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bui.

64:461^62. ♂.
Strumigenys (Cephaloxys) silvestriana Wheeler, 1928. Bol. Lab. Zool. Sci. Agr. Portici

22:27-28. ♂.
Strumigenys (Cephaloxys) foochowensis Wheeler, 1928. Bol. Lab. Zool. Sci. Agr. Portici

22:28-29. ♀.
Strumigenys (Cephaloxys) membranifera var. marioni Wheeler, 1933. Hawaii. Ent. Soc.,

Proc. 8:276. ♂.




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Strumigenys (Cephaloxys) membranifera var. vnlliamsi Wheeler, 1933. Hawaii. Ent. Soc,
Proc. 8:275-277. ♂, ♀.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:829, 856. —Brown, 1948. Amer.
Ent. Soc, Trans. 74:114. —Brown, 1949. Mushi 20:7, 22. —Wheeler and Wheeler, 1954.
Psyche 61:145 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:35 (Polynesia).

Biology: Weber, 1952. Amer. Mus. Novitates 1554:7. —Wilson, 1953. Ent. Soc. Amer., Ann.
46:483-485. —Brown, 1954. Harvard Univ., Mus. Comp. ZooL, Bui. 112:6.

Genus QUADRISTRUMA Brown

Qvxidristruma Brown, 1949. Amer. Ent. Soc, Trans. 75:47.
Type-species: Epitritus emmae Emery. Orig. desig.

The single species in the United States is an introduction.

Quadristruma emmae (Emery)
Fla. (homestead); Pantropical. Ecology: Most commonly found under objects

on sea beaches or in debris from forests or agricultural areas near the sea. Introduced;

spread to many tropical regions of the world by commerce, possibly originating from

Africa
Eiptritus emmae Emery, 1890. Soc. Ent. Ital., Bol. 22:70. ♂.
Epitritus clypeatus Szabo, 1909. Arch. Zool. (Budapest) 1(7):1. ♂.
Epitritus clypeatus var. malesiana Forel, 1913. Zool. Jahrb., Abt. f. System. 36:83-84. ♂.

9.
Epitritus wheeleri Donisthorpe, 1916. Ent. Rec. 28:121. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:149. —Smith, 1936. Puerto Rico
Univ., Jour. Agr. 20:829, 858. —Brown, 1949. Amer. Ent. Soc, Trans. 75:48-50 (worker,
female). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:42 (Polynesia).

Biology: Weber, 1934. Rev. de Ent. 4:51. —Brown, 1949. Mushi 20:21. —Brown, 1954.

Harvard Univ., Mus. Comp. Zool., BuL 112:6. —Brown and Wilson, 1960 (1959). Quart. Rev.
Biol. 34:281. — Kempf, 1961. Studia Ent. 4:516.

Tribe BASICEROTINI

Revision: Brown and Kempf, 1960. Studia Ent. 3:161-250 (world). —Taylor, 1968. Austral.
Jour. Zool. 16:333-348 (Indo-Australian).

Taxonomy: Wheeler and Wheeler, 1954. Psyche 61:111-145 (larvae).

Genus EURHOPALOTHRIX Brown and Kempf

Eurhopalothrix Brown and Kempf, 1960. Studia Ent. (n. s.) 3:202-203.

Type-species: Rhopalothrix Bolaui Mayr. Desig. by Brovvm and Kempf, 1961.

Taxonomy: Brown and Kempf, 1961 (1960). Psyche 67:44. —Wheeler and Wheeler, 1973.
Pan-Pacific Ent. 49:207 (larvae).

Eurhopalothrix floridana Brown and Kempf
Fla. (Highlands Hammock, Highlands Co.). Ecology: Collected in
leaf litter.
Eurhopalothrix floridana Brown and Kempf, 1960. Studia Ent. (n. s.) 3:207-208. ♂.

Tribe ATTINI

Taxonomy: Wheeler, 1948. Amer. Midland Nat. 40:664-689 (larvae). —Weber, 1970. Ent. Soc.
Wash., Proc 72:414-415 (northern extent of tribe). —Wheeler and Wheeler, 1974. Ent. Soc.
Wash., Proc 76:76-81 (larvae).

Biology: Weber, 1962. Amer. Mus. Nat. Hist., Bui. 71:45-49 (insect gardeners). —Weber,
1966. Science 153:587-604 (general). — Moser, 1967. Amer. Mus. Nat. Hist., Nat. Hist.
76:32-35 (trail of the leafcutters). —Weber, 1972. Amer. Scientist 60:448-456 (general).
— Weber, 1972. Amer. Phil. Soc, Mem. 92, 146 pp. (gardening ants, the attines).




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Morphology: Blum, Moser, and Cordero, 1964. Psyche 71:1-7 (source and specificity of odor
trail substances). — Moser, 1964. Science 143:1048-1049 (inquiline roach responds to trail
making substance). — Markl, 1970. Z. Vergl. Physiol. 69:2-37 (communication by
stridulatory signals). — Crewe and Blum, 1972. Jour. Ins. Physiol. 18:31-42 (alarm
pheromones, their phylogenetic significance).

Genus CYPHOMYRMEX Mayr

Cyphomyrmex Mayr, 1862. Zool.-Bot. CJesell. Wien, Verh. 12:690.

Type-species: Cyphomyrmex minutus Mayr. Monotypic.
Cyphomyrmex subg. Cyphomannia Weber, 1938. Rev. de Ent. 9:183.

Type-species: Cyphomyrmex {Cyphomannia) laexngatus Weber. Orig. desig.

A rather large Neotropical genus with extensions into the southern United States. These ants
form small colonies, usually in the soil, and live on fungi which they cultivate from a substratum
of vegetable matter of feces of certain insects.

Revision: Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:555-562. —Wheeler, 1907. Amer. Mus.
Nat. Hist., Bui. 23:670, 719-728, 765-773. —Weber, 1940. Rev. de Ent. 11:406-427. — Kempf,
1964. Studia Ent. 7:1^14 (strigatiis group). —Kempf, 1965. Studia Ent. 8:161-200 (rimosiis
group).

Taxonomy: Wheeler, 1948. Amer. Midland Nat. 40:668-670 (larvae). —Weber, 1966. Ent.
News 77:166-168.

Biology: Weber, 1955. Science 121:109. —Weber, 1957. Ecology 38: 480-494 (cultivation of

fungus).

Cyphomyrmex rimosus (Spinola)
Fla., Ala., Miss., La., Tex., Ariz., Calif.; W. Indies and Mexico s. to

Argentina. Ecology: Nests are in soil, under bark of rotten wood, or in humus around
roots; cultures a fungus in form of a yeast. A puzzling and variable complex for which
many subspecies have been proposed; see Kempf (1965) for names to be considered.

Cryptocerus '^ rimosus Spinola, 1853. Accad. Sci. Torino, Mem. (2) 13: 65. ♀, ♂.

Meranoplus difformis Smith, 1858. Cat. Hym. Brit. Mus. v. 6, p. 195. ♂.

Cyphomyrmex minutus Mayr, 1862. Zool.-Bot. Gtesell. Wien, Verh. 12:691. ♀.

Cyphomyrmex steinheili Forel, 1884. Soc. Vaud. des Sci. Nat., Bui. 20:368. ♂.

Cyphom,yrmex rimosus var. comxilensis Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:719.
5, 9, 6.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:719-712. —Weber, 1940. Rev. de
Ent. 11:411. —Weber, 1941. Rev. de Ent. 12:116-119. —Wheeler, 1948. Amer. Midland Nat.
40:668 (larva). —Weber, 1958. Ent. Soc. Wash., Proc. 60:259-260. —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62:28 Oarva). —Kempf, 1965. studia Ent. 8:162, 198.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:106. —Weber, 1945. Rev. de Ent.
16:5-14. —Weber, 1947. Bol. de Ent. Venezolana 6:144. —Weber, 1955. Wash. Acad. Sci.,
Jour. 45:275-281. —Weber, 1972. Amer. Phil. Soc., Mem. 92, pp. 8, 26, 31-32, 34, 39-42, 51,
60, 71-72, 94-95, 98-99, 102, 105-108, 110, 117.

Cyphomyrmex wheeleri
***authority mismatch
ForeL Tex., Calif.; Mexico. Ecology: Nests in the soil beneath stones in very arid
habitats.
Cyphomyrmex wheeleri Forel, 1900. Schweiz. Ent. (Jesell. Mitt. 10:282. 9,9.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:725-726 (each caste). —Weber,
1940. Rev. de Ent. 11:409. —Kempf, 1965. Studia Ent. 8:167-172.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40:74. —Weber, 1972. Amer. PhiL Soc.,
Mem. 92, pp. 26, 93. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 97-98.

Genus MYCETOSORITIS Wheeler

Atta subg. Mycetosoritis Wheeler, 1907. Amer. Mus. Nat. Hist, BuL 23:714.
Type-species: Atta (Mycetosoritis) hartmanni Wheeler. Monotypic

A small genus found only in the United States and Brazil.




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Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104:317-318. — Kempf,
1964. Studia Ent. 7:1.

Mycetosoritis hartmanni (Wheeler)
La., Tex. Ecology: Forms small colonies in soil; food consists of a fungus
grown upon a substratum composed of flower anthers.
Atta (Mycetosoritis) hartmanni Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:714. ♂, ♀.

6.

Taxonomy: Wheeler and Wheeler, 1974. Ent. Soc. Wash., Proc. 76:77-79 (larva).

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:761-765. —Wheeler, 1910. Ants, pp.
334-335.

Genus TRACHYMYRMEX Forel

Atta subg. Trachymyrmex Forel, 1893. Soc. Ent. de Belg., Ann. 37:600.

Tyi)e-species: Atta septentrionalis McCook. Desig. by Wheeler, 1911.

A Neartcic and Neotropical genus found from New York south to Argentina, and the largest
Genus of attine ants. In the United States, most forms are recorded from the more humid areas,
though a few occur in arid sections. Nests are in the soU. The fungi on which the ants feed is
grown on a substratum of plant particles and insect excrement.

Revision: Wheeler, 1907. Amer. Mus. Nat. Hist, Bui. 23:706-714, 746-760. —Wheeler, 1911. N.
Y. Ent. Soc., Jour. 19:245-255.

Taxonomy: Weber, 1945. Rev. de Ent. 16:44. —Wheeler, 1948. Amer. Midland Nat. 40:673-674
(larvae).

Biology: Weber, 1955. Science 121:109. —Weber, 1960. Ent. News 71:1-6 (comparison of
worker and female behavior).

arizonensis (Wheeler). Ariz. (Huachuca Mtns.). Ecology: Found in arid, stony canyons, 5000 to
6000 ft.
Atta (Trachymyrmex) arizonensis Wheeler, 1907. Amer. Mus. Nat. Hist., BuL 23:710. ♀, ♂.

Taxonomy: Wheeler, 1911. N. Y. Ent. Soc., Jour. 19:250 (worker).

Biology: Wheeler, 1911. Psyche 18:93-98. — Essig, 1926. Ins. of West. No. Amer., p. 862.
—Weber, 1972. Amer. PhiL Soc., Mem. 92, p. 93.
desertorum (Wheeler). Ariz. (nr. Tucson). Ecology: Nests were found in hard, pebbly, desert
soil.
Atta (Trachymyrmex) desertorum Wheeler, 1911. Psyche 18:98-100. ♂.

Taxonomy: Wheeler, 1911. N. Y. Ent. Soc., Jour. 19:249.

Biology: Wheeler, 1911. Psyche 18:100-101. —Essig, 1926. Ins. of West. No. Amer., p. 862.
jamaicensis (Andre). Fla. (Dania); W. Indies. Possibly introduced.

Atta (Acromyrmex) jamaicensis Andre, 1893. Rev. Ent. Caen 12:149. 2.

Trachymyrmex sharpi Forel, 1893. Ent. Soc. London, Trans., p. 372. ♂.

Atta (Trachymyrmex) maritima Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:107. ♂.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:860. —Weber, 1967. Ent. News
78:107-109.

Biology: Weber, 1972. Amer. Phil. See., Mem. 92, pp. 102, 104 (fungi).
nogalensis Byars. Ariz. (Nogales).

Trachymyrmex nogalensis Byars, 1951. Ent. Soc. Wash., Proc. 53:109-111. ♂, ♀.
septentrionalis (McCook). N. Y. (L. I., Staten Is.) s. to Fla., w. to 111., La, Tex. Ecology: Nests
are in soil, usually with a small semicircular crater. Of some economic importance
because of the habit of cutting leaves from domesticated plants. The New York records
are the northernmost for an attine ant.
Oecodoma virginiana Buckley, 1867. Ent. Soc. Phila., Proc. 6:346. 5 . Placement of this

species is questionable.
Atta septentrionalis McCook, 1880. Acad. Nat. Sci. Phila., Proc. 32:359. ♂.
Atta (Trachymyrmex) septentrionalis var. obscurior Wheeler, 1907. Amer. Mus. Nat. Hist.,
Bui. 23:706. 9, 9, cJ.




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Atta (Trachymyrmex) septentnonalis var. vertebrata Wheeler, 1911. N. Y. Ent. Soc, Jour.

19:246, 250. ♂, ♀.
Atta (Trachymyrmex) septentrionalis obscurior var. sem.inole Wheeler, 1911. N. Y. Ent.

Soc, Jour. 19:247, 250. $, 9, cJ.
Atta (Trachymyrmex) septentrionalis obscurior var. crystallina Wheeler, 1911. N. Y. Ent.

Soc, Jour. 19:247,250. ♂, ♀.
Atta (Trachymyrmex) septentrionalis obscurior var. irrorata Wheeler, 1911. N. Y. Ent.

Soc, Jour. 19:247,250. ♂.

Taxonomy: Forel, 1884. Soc. Vaud. des Sci. Nat., Bui. 20:91. —Smith, 1924. Ent. News 35:52.
—Cole, 1940. Amer. Midland Nat. 24:59. —Wheeler, 1948. Amer. Midland Nat. 40:673
(larva). —Weber, 1958. Ent. News 69:52-53.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21:374, 386-387. —Wheeler, 1907. Amer.
Mus. Nat. Hist., Bui. 23:680-681, 746-753. —Wheeler, 1911. Psyche 18:95-97. —Wheeler,
1911. N. Y. Ent. Soc, Jour. 19:250-255. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:273, 292,
305. —Cole, 1939. Lloydia 2:153-160. —Cole, 1951 (1950). Ent. Soc. Amer., Ann. 43:499-500.
—Weber, 1956. Ecology 37:150-161. —Weber, 1966. Ent. News 77:241. — Moser, 1964.
Science 143:1048-1049 (inquiline roach responds to trail-making substance). — Weber, 1972.
Amer. Phil. Soc, Mem. 92, pp. 3, 13-15, 25-28, 32-33, 35-36, 40-44, 46, 51, 54-55, 62, 70, 99,
103-104, 106-107, 112, 117.
smithi neomexicanus Cole. N. Mex., Ariz. Ecology: Nests found in sand. T. smithi sm,ithi
Buren occurs in Mexico.
Trachymyrmex smithi neomexicamis Cole, 1952. Tenn. Acad. Sci., Jour. 27:159-162. ♂.

Taxonomy: Cole, 1953. Tenn. Acad. Sci., Jour. 28:300-301 (female, male).
turrifex caroli (Wheeler). Tex. (Huntsville).

Atta (Trachymyrmex) turrifex caroli Wheeler, 1911. N. Y. Ent. Soc, Jour. 19:248. ♂.
turrifex turrifex (Wheeler). La., Tex.; Mexico. Ecology: Nests in soil.

Atta (Trachymyrmex) turrifex Wheeler, 1903. Psyche 10:100. ♂, ♀.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat., Hist., Bui. 23:709-710. —Wheeler, 1911. N. Y.
Ent. Soc, Jour. 19:249.

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:753-759. —Wheeler, 1911. Psyche
18:95-97.

Genus ACROMYRMEX Mayr

Atta subg. Acromyrmex Mayr, 1865. Reise d. Novara, Zool. v. 1, Hym., p. 83.

Type-species: Formica hystrix Latreille. Monotypic
Atta subg. Moellerius Forel, 1893. Soc. Ent. de Belg., Ann. 37:589.

Type-species: Atta (Acromyrmex) landolti Forel. Desig. by Wheeler, 1911.
Pseudoatta Gallardo, 1916. Buenos Aires Mus. Nac de Hist. Nat., An. 28:320. Syn.
uncertain.

Type-species: Pseudoatta argentina Gallardo. Monotypic.

A Neotropical genus with a slight extension into southwestern United States. These are
mostly grassland and semidesert fungus-growing ants which usually collect grass. They are com-
parable to Atta in biology and economic importance but colonies are apparently not so large and
the leaf -cutting habits not so extensive. The species listed below have previously been placed in
the subgenus Moellerius.

Revision: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:669-700, 703-706, 743-746. — Santschi,
1925. Rev. Suisse de Zool. 31:386-398.

Taxonomy: Weber, 1945. Rev. de Ent. 16:61-62. —Wheeler, 1948. Amer. Midland Nat.
40:674-676 (larvae). — Brown, 1973. In Meggers, et al.. Tropical Forest Ecosystems in Afr.
and S. Amer., pp. 178-185 (generic syn.).

Biology: Weber, 1967. Ent. Soc. Amer., Ann. 78:107-109 (growth of young colonies in first
year).




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Acromyrmex versicolor
***authority mismatch
chisosensis (Wheeler). Tex. (Mtns. of Big Bend area).

Atta (Moellerius) versicolor chisosensis Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:705.
9.

Acromyrmex versicolor
***authority mismatch
versicolor (Pergande). Ariz., Calif.; Mexico. Ecology: Nests are in soil in deserts;
collects pieces of leaves from bushes and herbs for a media on which to grow fungus.
Atta versicolor Pergande, 1893. Calif. Acad. Sci., Proc. 4:31. ♂.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:703-705 (each caste).

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:743-746. —Wheeler, 1911. Psyche
18:101. —Wheeler, 1917. Psyche 24:179-180. —Weber, 1972. Amer. Phil. Soc., Mem. 92, pp.
22, 26, 30, 32.

Genus ATTA Fabricius

Atta Fabricius, 1804. Systema Piezatorum, p. 421.

Type-species: Formica cephalotes Linnaeus. Desig. by Wheeler, 1911.
Oecodoma Latreille, 1818. Nouv. Diet. Hist. Nat. 23:223.

Type-species: Formica cephalotes Linnaeus. Desig. by Shuckard, 1840.
Archeatta Goncalves, 1942. Soc. BrasU. de Agron., Bol. 5:342.

Type-species: Oecodoma meocicana Smith. Orig. desig.
Atta subg. Neoatta Goncalves, 1942. Soc. Brasil. de Agron., Bol. 5:334.

Type-species: Formica sexdens Linnaeus. Orig. desig.
Atta subg. Palaeatta Borgmeier, 1950. Inst. Oswaldo Cruz, Mem. 48:244, 270.

Type-species: Atta bisphaerica Forel. Orig. desig.
Atta subg. Epiatta Borgmeier, 1950. Inst. Oswaldo Cruz, Mem. 48:246, 272.

Type-species: Atta la^evigata Smith. Orig. desig.

A Neotropical genus with an extension into southern United States. The nests, which are deep
in the soU and usually have many lateral and vertical entrances, are often extremely large and
contain numerous individuals. Workers defoliate both wild and domesticated plants, including
trees, and they feed on a fungus grown from a substratum composed of macerated leaves and
other vegetable matter.

Revision: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:669-703, 729-742. —Goncalves, 1942.
Soc. Brasil. de Agron., Bol. 5:333-358. —Borgmeier, 1959. Studia Ent. 2:321-390.

Taxonomy: Wheeler, 1948. Amer. Midland Nat. 40:676-683 (larvae). —Borgmeier, 1950. Inst.
Oswaldo Cruz, Mem. 48:239-263. —Weber, 1958. Ent. News 69:7-13. —Smith, 1963. Ent.
Soc. Wash., Proc. 65:299-302 (key to spp. of U. S. and Mexico; bioL notes). —Weber, 1968.
Ent. Soc. Wash., Proc. 70:348-350 (Panama species).

Biology: Weber, 1962. Amer. Mus. Nat. Hist., Nat. Hist. 61:45-51.

Morphology: Blum, et al., 1968. Comp. Biochem. Physiol. 26: 291-299 (alkanones and terpenes
in the mandibular glands of Atta species).

Atta mexicana (Smith)
Ariz. (Organ Pipe Cactus Natl. Mon.); Mexico s. to Honduras. Ecology:

Nests are in a variety of ecological situations and their fungus gardens are grown on a
wide variety of vegetable substances.
Oecodoma mexicana Smith, 1858. Cat. Hym. Brit. Mus. 6:185. ♀, ♂.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:702-703. —Goncalves, 1942. Soc.

Brasil. de Agron., Bol. 5:335, 337, 343. — Byars, 1949. Jour. Econ. Ent. 42:545 (Ariz, record;

soldier). —Borgmeier, 1950. Inst. Oswaldo Cruz, Mem. 48:260-261. —Borgmeier, 1959.

Studia Ent. 2:351-353 (each caste). —Smith, 1963. Ent. Soc. Wash., Proc. 65:299-302.

Atta texana (Buckley)
La., Tex.; e. Mexico. Ecology: Nests are in well-drained sand or loamy soils.
The interior of the nest may reach a depth of 15 to 20 ft. and contains innumerable
interconnected chambers. It is connected to the exterior by numerous holes which are
surrounded by crater-shaped piles of dirt. In a large nest there may be 1,000 entrance
holes occupying an area of 4,500 square feet. The ants grow a fungus primarily on
macerated leaves. An economicaUy important pest; invades houses; cuts leaves from
domesticated plants; steals seeds; builds unsightly nests; damages roads, walks, stock, or




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equipment by cave-ins of the nest; and can inflict painful bites. Texas leafcutting ant;
also known locally as the town ant, night ant, cut ant, parasol ant, pack ant, or fungus
ant.
Myrmica (Atta) texana Buckley, 1860. Acad. Nat. Sci. Phila., Proc. 12:233. 5, 9, S.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:700-703. — Goncalves, 1942. Soc.
Brasil. de Agron., Bol. 5:335, 337, 343. — Byars, 1949. Jour. Econ. Ent. 42:545. — Borgmeier,
1950. Inst. Oswaldo Cruz, Mem. 48:260-261. —Borgmeier, 1959. Studia Ent. 2:356-358.
—Smith, 1963. Ent. Soc. Wash., Proc. 65:299-301. —Wheeler and Wheeler, 1974. Ent. Soc.
Wash., Proc. 76:79-81 (larva).

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bui. 23:729-742. —Hunter, 1912. U. S. Dept.
Agr., Bur. Ent. Cir. 148:1^. ^ones, 1917. Jour. Econ. Ent. 10:561. —Snyder, 1937. La.
Conserv. Rev., pp. 14-17. —Walter, Seaton, and Mathewson, 1938. U. S. Dept. Agr. Cir.
494:1-18. — Metcalf and Flint, 1939. Destructive and Useful Insects, p. 771. —Smith, 1939.
South. Forest Expt. Sta., Occas. Papers 84:2-6. -Johnston, 1944. Jour. Forestry 42:130-132.
—Bennett, 1958. U. S. Dept. Agr., Forest Pest Leaflet 23:1-4 (rev. 1967). — Warter, Moser,
and Blum, 1962. La. Acad. Sci., Proc. 25:42-46 (correlation between night hawk feeding and
nuptial flights). — Moser, 1962. Forests and People 12:40-41 (nesting habits). — Moser,
1963. Ent. Soc. Amer., Ann. 56:286-291 (contents and structure of nest in summer).
— Moser, 1964. Science 143:1048-1049 (inquiline roach responds to trail-making substance).
— Moser, 1967. Insectes Sociaux 14:295-312 (mating activities). — Moser, 1967. Amer. Mus.
Nat. Hist., Nat. Hist. 76:33-35. —Weber, 1972. Amer. Phil. Soc., Mem. 92, pp. 3, 24, 30-33,
35, 37-38, 65, 81, 84.

Morphology: Moser and Blum, 1963. Science 140:1228 (trail marking substance, source and
potency). — Moser, Brownlee, and Silverstein, 1968. Jour. Ins. Physiol. 14:529-535 (alarm
pheromones). — Hermann, Moser, and Hunt, 1970. Ent. Soc. Amer., Ann. 63:1152-1158
(poison apparatus, morphological and behavioral changes). — Moser and Silverstein, 1967.
Nature 215:206-207 (volatility of trail marking substance). — Tumlinson, et al., 1971.
Nature 234: 348-349 (identification of trail pheromone). — Tumlinson, et al., 1972. Jour.
Insect. Physiol. 18: 809-814 (a volatile trail pheromone). —Riley, et al., 1974. Science 183:
760-762 (biological responses to its alarm pheromone).

Unplaced Taxa of Myrmicinae

Pheidole buckleyi Smith, 1951. In Muesebeck, et al, U. S. Dept. Agr., Agr. Monog. 2:806. N.

name for Atta pennsylvanica Buckley.
Crematogaster (Acrocoelia) sanguinea coachellai Enzmann, 1946. N. Y. Ent. Soc., Jour. 54:95.

5 . In same paper, fig. 3 of the petiole is given as that of Crematogaster {Acrocoelia)

lineolata var. coachellai.
Myrmica corrugata Say, 1836. Boston Jour. Nat. Hist. 1:291. ♀, ♂.
Myrmica dimidiata Say, 1836. Boston Jour. Nat. Hist. 1:293. 9 (?). N. Amer.
Myrmica (MonoTnarium (.')) diversa Buckley, 1867. Ent. Soc. Phila., Proc. 6:337. 5,9. Tex.
Myrmica inflecta Say, 1836. Boston Jour. Nat. Hist. 1:292. ♂.
Myrmica (Monomarium (.')) lineolata Buckley, 1867. Ent. Soc. Phila., Proc. 6:340. ♂, ♀. Preocc.

by Say, 1836. No locality.
Solenopsis madara Roger, 1863. Berlin. Ent. Ztschr. 7:200. ♂, ♀.

Myrmica (Monomarium (.')) montana Buckley, 1867. Ent. Soc. Phila., Proc. 6:339. 5. Tex.
Myrmica novaeboracensis Buckley, 1867. Ent. Soc. Phila., Proc. 6:337. 9. N. Y.
Myrmica opposita Say, 1836. Boston Jour. Nat. Hist. 1:292. 5, cJ. N. Amer.
Atta pennsylvanica Buckley, 1867. Ent. Soc. Phila., Proc. 6:345. 5, 1(. Pa. Previously placed in

Pheidole where it would be preocc. by Roger, 1863; see Pheidole buckleyi Smith.
Atta picea Buckley, 1867. Ent. Soc. Phila., Proc. 6:344. 5, V. Tex.
Oecodoma pilosa Buckley, 1867. Ent. Soc. Phila., Proc. 6:348. 5 . Tex.
Myrmica (Diplorhoptrum) scabrata Buckley, 1867. Ent. Soc. Phila., Proc. 6:343. 5 . Conn.
Myrmica (Atta) sublanuginosa Buckley, 1867. Ent. Soc. Phila., Proc. 6:343. 5 . Tex.
Oecodoma tardigrada Buckley, 1867. Ent. Soc. Phila., Proc. 6:349. 5, 9, S. Tex.
Pheidole pubiventris race timmi Forel, 1901. Mitt. Naturh. Mus. Hamburg 18:62. 5, 'U.

"Probably of Mexican origin - Dr. R. Timm."




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Subfamily DOLICHODERINAE

A small subfamily with most Nearctic representatives found in the southern portion of the
United States. Most species are small, drab-colored insects and are recognized by the single seg-
mented pedicel, lack of a constriction between the first and second gastric segments, and the
slitlike cloacal orifice.

Taxonomy: Starcke, 1933. Tijdschr. v. Ent. 76: XXVI-XXXII (larvae). —Wheeler and
Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 169-210 (larvae). —Wheeler and Wheeler, 1966.
Ent. Soc. Amer., Ann. 59: 726-732 (larvae). —Wheeler and Wheeler, 1974 (1973).
Pan-Pacific Ent. 49: 396-401 (larvae).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bui. 116: 453-464
(proventriculus; use as a generic character).

Tribe DOLICHODERINI

Genus DOLICHODERUS Lund

Brown (1973) listed Monads Roger, 1862, Diabolus Karavaiev, 1926, Diceratoclinea Wheeler,
1935, and Karawajewella Donisthorpe, 1944 as synonyms of Dolichoderus and Hypoclinea Mayr,
1855, and Acanthoclinea Wheeler, 1935 as provisional synonyms. Because of the doubt as to the
exact status of some of the genera, subgeneric groupings are used as in some previous North
American literature until the genus has been adequately studied.

Taxonomy: Brown, 1973. In Meggers, et al.. Tropical Forest Ecosystems in Afr. and S.
Amer., pp. 178-185 (generic syn.).

Genus DOLICHODERUS Subgenus DOLICHODERUS Lund

Dolichodenis Lund, 1831. Ann. des Sci. Nat, Zool. 23: 130.

Type-species: Formica attelahovdes Fabricius. Monotypic.

Not known to occur in the Nearctic Region.

Genus DOLICHODERUS Subgenus HYPOCLINEA Mayr

Hypoclinea Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5: 377.

Type-species: Formica quadrijmnctatxi Linnaeus. Desig. by Wheeler, 1911.

In North America, this subgenus is found only in the eastern half of the continent. Colonies
are smaU to moderately large and nests are constructed in the soil, in curled leaves and hollow
stems of plants, and in cartons attached to plants. Workers attend honeydew-excreting insects
and feed on small arthropods. Some forms emit a fluid which has a peculiar smoky or pungent
odor.

Revision: Mayr, 1870. Zool.-Bot. GeseU. Wien, Verh. 20: 953-960. —Mayr, 1886. Zool.-Bot.
Gesell. Wien, Verh. 36: 434-437. —Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 305-319.

Taxonomy: Brown, 1950. Wasmann Jour. Biol. 8: 249. —Wheeler and Wheeler, 1951. Ent. Soc.
Wash., Proc. 53: 172 (larvae). —Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59:
726-730 Oarvae).

Biology: Kannowski, 1959. Insectes Sociaux 6: 129-134 (flight comparisons of some spp.).
— Torossian, 1960. Insectes Sociaux 7: 383-393 (biology of D. qtuidripunctatus (L.)).

Morphology: Torossian, 1959. Insectes Sociaux 6: 369-374 (tropholactic and proctodeal
exchange in D. qtuidripunctatus (L.)).

Dolichoderus mariae Forel
Mass. to Ga., w. to Minn., 111., Okla., La. Ecology: Nests of the rather large

colonies are found in the soil, preferably sand, beneath tufts of grass or smaU bushes.
Dolichodenis Mariae Forel, 1884. Soc. Vaud. des Sci. Nat., Bui. 20: 349. ♂.
Dolichoderus mariae davisi Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 306, 308. ♂.
Dolichodenis (Hypoclinea) mariae var. blatchleyi Wheeler, 1917. Ind. Acad. Sci., Proc. 26:
462. ♂.




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Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 306. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bui. 22: 589 (worker, female). —Cole, 1940. Amer. Midland
Nat. 24: 60. —Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 726 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 316-319, 387-388. —Smith, 1924. Ent.
News 35: 81. —Talbot, 1956. Psyche 63: 134-139 (flight activities). — Kannowski, 1959.
Insectes Sociaux 6: 129-133.

Dolichoderus plagiatus (Mayr)
N. B., Ont. s. to Ga., Tenn., w. to N. Dak., 111. Ecology: Colonies are small and

nests are found in inconspicuous places such as under forest debris in the soil, in hollow

stems, and in curled-up leaves.
Hypoclinea plagiata Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 957, 960. ♂.
Dolichoderus borealis Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 408. ♂.
Dolichoderus plagiatus var. inomatus Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 306,

313. ♀.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 310 (each caste). —Wheeler, 1917.
Conn. State Geol. and Nat. Hist. Survey Bui. 22: 590. —Cole, 1940. Amer. Midland Nat. 24:
60. —Gregg, 1944. Ent. Soc. Amer., Ami. 37: 467. —Wheeler, and Wheeler, 1966. Ent. Soc.
Amer., Ann. 59: 727 (larva). — Francoeur and Beique, 1966. Canad. Ent. 98: 142
(Provancher types).

Biology: AVheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 306, 315-316, 388. —Wheeler, 1908.
Amer. Mus. Nat. Hist., Bui. 24: 621-622. —Wesson and Wesson, 1940. Amer. Midland Nat.
24: 90, 99. —Kannowski, 1959. Insectes Sociaux 6: 133. —Wheeler and Wheeler, 1963. Ants
of N. Dak, pp. 149-151. —Kannowski, 1967. Ent. Soc. Amer., Ami. 60: 1246-1252
(population studies).

Dolichoderus pustulatus Mayr
N. S. s. to Fla., w. to 111., Okla., Tex. Ecology: Colonies are small and nests
are found under piles of detritus or in a hard, thin carton shell above ground about
blades of a tuft of grass.

Dolichoderus frmtulatiis Mayr, 1886. Zool.-Bot. Gesell. Wien., Verh. 36: 435-436. ♂, ♀.

Dolichoderus plagiatiis var. beutenmuelleri Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:
304. ♀.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 306, 313, 388. —Smith, 1924. Ent.
News 35: 82. —Smith, 1931. Ent. News 42: 22. —Wheeler and Wheeler, 1966. Ent. Soc.
Amer., Ann. 59: 727-728 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 315-316, 388. —Wheeler, 1917. Ind.

Acad. Sci., Proc. 26: 462. —Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 12. —Wesson and

Wesson, 1940. Amer. Midland Nat. 24: 89, 99. — Kannowski, 1959. Insectes Sociaux 6:

133-134, 155. —Kannowski, 1967. Ent. Soc. Amer., Ann. 60: 1246-1252 (population studies).

taschenberg^ (Mayr). N. S., Ont. s. to S. C, w. to Man., N. Dak., S. Dak., La. Ecology: Nests of

rather large colonies are constructed in the soil in wooded areas or the edge of woods

and usually have a low moimd of thatch (grass, twigs, needles) over the entrance hole.

Workers commonly attend honeydew-excreting insects.
Hypoclinea Taschenbergi Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 53:

498. ♀.
Dolichoderus taschenbergi var. gagates Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306,

310. ♀. Preocc. by Emery, 1890.
Dolichoderus {Hypoclinea) taschenbergi var. aterrimus Wheeler, 1915. Amer. Mus. Nat.

Hist., Bul. 34: 417. N. name for gagates Wheeler.
Dolichoderus tachenbergi var. wheeleriella Forel, 1916. Rev. Suisse de Zool. 24: 458. N.

name for gagates Wheeler.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306, 309-310. —Smith, 1924. Ent.
News 35: 82. —Cole, 1940. Amer. Midland Nat. 24: 60. —Wheeler and Wheeler, 1966. Ent.
Soc. Amer., Ann. 59: 728 Garva).

Biology: Wheeler, 1915. Psyche 22: 306. —Smith, 1924. Ent. News 35: 81-82. — Logier, 1923.
Canad. Ent. 55: 247. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 292-293. —Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 99. —Kannowski, 1959. Ent. Soc. Amer., Ann. 52:




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755-760 (flight activities). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 151-154.
— Bradley and Hinks, 1968. Canad. Ent. 100: 40-50 (attending aphids on jack pine).
— Bradley, 1972. Canad. Ent. 104: 245-249 (transplanting colonies in jack pine stands in
Man.). —Bums, 1973. Canad. Ent. 105: 97-104 (foraging and tending behavior).

Tribe TAPINOMINI

Genus LIOMETOPUM Mayr

Liometopum Mayr, 1861. Die Europaischen Formiciden, pp. 25, 38.
Type-species: Formica microcephala Panzer. Monotypic.

A Holarctic genus found only in the western United States in the Nearctic region. The Nearc-
tic forms nest in the soil beneath cover or under bark or in crevices in trees and the nest cham-
bers are usually subdivided by a network of paperlike material. Colonies are often populous.
Workers forage in files sometimes several hundred feet long and commonly attend coccids and
aphids. They are pugnacious and eject a repellent secretion with a strong and disagreeable odor
like that of butyric acid.

Revision: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 321-333.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 172, 181 (larvae).

Liometopum apiculatum Mayr
W. Tex., Colo., N. Mex., Ariz.; Mexico. Ecology: Usually in foothill areas at
elevations of 4000 to 7000 feet.
Liometopum apiculatum Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 961. ♂.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 331 (female). —Wheeler, 1905.
Amer. Mus. Nat. Hist., Bui. 21: 322-324 (each caste). —Wheeler and Wheeler, 1951. Ent.
Soc. Wash., Proc. 53: 181 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 322-324. —Wheeler, 1917. Psyche 24:
177-178. —Gregg, 1963. Univ. Colo. Studies, Biol. Ser. 11, pp. 1-6. —Gregg, 1963. Ants of
Colo., pp. 438-441. —Van Pelt, 1971. Ent. Soc. Amer., Ann. 64: 1186 (trophobiosis and
feeding habits).

Liometopum occidentale
***authority mismatch
luctuosum Wheeler. Wyo., Colo., w. Tex., w. to Ariz., Calif. Ecology: At higher
elevations, 4000 to 8000 feet.
Liometopum- apiculatum, luctuosum. Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 325. ♀.

Taxonomy: Forel, 1914. Deut. Ent. Ztschr. 6: 619 (male). —Cole, 1942. Amer. Midland Nat.
28: 371.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 332. — Mallis, 1941. South Calif. Acad.
Sci., Bui. 40: 75. —Gregg, 1963. Ants of Colo., pp. 441-443. —Cole, 1966. Brigham Young
Univ., Sci. Bui., Biol. Ser. 7: 18 (Nevada Test Site). —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 100-101.

Liometopum occidentale
***authority mismatch
occidentale Emery. Oreg., Calif.; Mexico. Ecology: Foimd at lower elevations, sea
level to 4000 feet.
Liometopum, microcephalum var. occidentale Emery, 1895. Zool. Jahrb., Abt. f. System. 8:
330. $, 9.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 324-325. — Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 8, 17.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 522. —Mallis, 1941. South Calif.
Acad. Sci., Bui. 40: 75. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 100.

Genus IRIDOMYRMEX Mayr

Iridomyrmex Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 702.

Type-species: Formica detecta Smith. Desig. by Bingham, 1903.
Tapinoma subg. Dolerom.yrma Forel, 1907. Mus. Nat. Hungarici, Ann. 5: 28.

Type-species: Tapinoma {Doleromyrma) darwinianum, Forel. Monotypic.
Anonychamyrma Donisthorpe, 1946. Ann. and Mag. Nat. Hist. 13: 588. Syn. uncertain.

Type-species: Anonychomyrma myrmex Donisthorpe. Monotypic.




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Three of the four species of this cosmopolitan genus found in North America have been in-
troduced. Most species nest in the soil and some are found in buildings. Workers collect
honeydew but are also entomophagous.

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 953-960. —Smith, 1929. Jour. Econ.
Ent. 22: 241-243.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 172, 184 (larvae).
—Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 729 (larvae).

Iridomyrmex glaber (Mayr)
Fla. (Winter Park); Australian and Oriental regions. Ecology: Florida specimens
collected from a stump of queen palm, Arecastruni romanzqffianum. Probably
introduced. These specimens are tentatively referred to glaber since the taxonomy of

Iridomyrmex glaber
***authority mismatch
and related species in the Old World are unstudied.
Hypoclinea glabra Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 705. 5, d.

Taxonomy: Brovtm, 1958. Acta Hym. 1:40-41 (also biological notes; New Zealand; Australia).
—Wheeler and Wheeler, 1974 (1973). Pan-Pacific Ent. 49: 398 (larva, from Australian
specimens).

Iridomyrmex humilis (Mayr)
Md. to Fla. w. to 111., Tex., Ariz., Calif.; Mexico, S. Amer.; Europe; S. Africa;
Hawaii; Australia. Ecology: Nests in soil, rotting wood or debris and lives in large
colonies that contain many females. Workers forage in files and tend
honeydew-excreting insects though they also feed on other sweet substances. A most
persistent and troublesome house-infesting ant; also known to steal seeds from
seedbeds, drive poultry from their nests, kiU newly hatched chickens, foster
honeydew-excreting insects, disrupt bee hives, and gnaw into ripened fruits. Introduced,
native to S. Amer. and spread to the U. S. and other parts of the world by commerce;
probably introduced first into New Orleans on coffee ships from Brazil sometime prior
to 1891. Argentine ant.
Hypoclinea {Iridomyrmex) humilis Mayr, 1868. Soc. Nat. Modena, Ann. 3: 164. ♀.

Taxonomy: Wheeler, 1913. In Newell and Barber, U. S. Dept. Agr. Bur. Ent. Bui. 122: 27-30
(each caste). —Smith, 1929. Jour. Econ. Ent. 22: 241. —Wheeler and Wheeler, 1951. Ent.
Soc. Wash., Proc. 53: 186 (larva). —Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59:
729 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 79-80 (Polynesia).

Biology: Newell, 1909. Jour. Econ. Ent. 2: 174-192 (life history). —Newell and Barber, 1913.
U. S. Dept. Agr. Bur. Ent., Bui. 122: 1-98. — Gallardo, 1915. Buenos Aires Mus. Nac. de
Hist. Nat., An. 27: 23-25. —Barber, 1920. U. S. Dept. Agr. Farmers' Bui. 1101: 1-11.
— Woglum and Borden, 1921. U. S. Dept. Agr. Bui. 965: 1-43. — Hamed and Smith, 1922.
Jour. Econ. Ent. 15: 261-264. — Essig, 1926. Ins. of West. N. Amer., pp. 865-866. —Smith,
1936. U. S. Dept. Agr. Cir. 387: 1-39. — Mallis, 1942. Sci. Monthly 55: 536-545 (half a
century with the Argentine ant). —Flanders, 1943. Calif. Citrograph 28: 117, 128, 137
(Argentine ant versus parasites of black scale). — McCluskey, 1958. Science 128: 536-537
(daily rhythm in male Argentine ant). — Skaife, 1962. The Study of Ants, p. 12. — Pavan,
1963. Pavia Univ., Sympos. (Jenet. et Biol. Ital. 12: 122-131. — HasWns and Haskins, 1965.
Ecology 46: 737 (extension of range in Bermuda at expense of Pheidole megacephala (F.)).
—Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 52-54 (economic importance).
— Markin, 1968. Kans. Ent. Soc., Jour. 41: 511-516 (study in citrus groves in Calif.).
— Markin and McCoy, 1968. Ent. Soc. Amer., Ann. 61: 505-509 (a nematode in pharyngeal
glands). — Markin, 1969. Insectes Sociaux 16: 159-190 (food distribution in laboratory
colonies). — Dechene, 1970. Wasmann Jour. BioL 28: 175-184 (behavioral patterns).
— Fluker and Beardsley, 1970. Ent. Soc. Amer., Ann. 63: 1290-1296 (ecological notes, in
Hawaii). — Markin, 1970. Jour. Econ. Ent. 63: 740-744 (foraging behavior in Calif, citrus
groves). —Markin, 1970. Insectes Sociaux 17: 127-158. —Markin, 1970. Ent. Soc. Amer.,
Ann. 63: 1238-1242 (seasonal life cycle in S. Calif.). — Erickson, 1971. Psyche 78: 257-266
(displacement of native ant spp. by Argentine ant).

Morphology: Pavan, 1950 (1948). 8th Intemat. Cong. Ent., Proc., Stockholm, pp. 863-865
(physiology). —Pavan, 1955. Soc. Ital. di Sci. Nat. Atti 94: 379-477. —Weber, 1961. Ent.
Soc. Wash., Proc. 63: 218 (use of poison). — Torossian, 1961. Insectes Sociaux 8: 189-191.
—Nachtwey, 1963. Insectes Sociaux 10: 50-53.




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iniquus nigellus Emery. Mass., 111., other scattered localities mostly in northeastern U. S.;
Central Amer. Ecology: Found in greenhouses in U. S. Introduced. /. iniquus iniquus
Mayr occurs in Central and S. Amer. and W. Indies.
Iridoniyrmex iniquus var. nigellus Emery, 1890. Soc. Ent. Ital., Bol. 22: 56. ♂.

Biology: Wheeler, 1929. Psyche 36: 89-90. —Smith, 1929. Jour. Econ. Ent. 22: 241-242.

Iridomyrmex pruinosus
***authority mismatch
analis (Andre). N. Dak., Kans., Okla., Tex. w. to s. Idaho, Calif.; Mexico, Guatemala.
Ecology: Nests are in soil either under objects or in exposed situations surmoimted by a
craterlike mound.
Tapinoma anale Andre, 1893. Rev. Ent. de France 12: 148. ♂.
Iridoniyrmex pruinosus var. testaceus Cole, 1936. Ent. News 47: 121. ♂, ♀.

Taxonomy: Smith, 1929. Jour. Econ. Ent. 22: 241. — Eckert and Mallis, 1937. Calif. Agr. Expt.
Sta. Cir. 342: 8. —Cole, 1942. Amer. Midland Nat. 28: 372-373 (also biological notes).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 342. —Smith, 1924. Ent. News 35: 83.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31: 294, 305. —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 154-155. —Gregg, 1963. Ants of Colo., pp. 436-438. —Cole, 1966. Brigham
Young Univ., Sci. Bui., Biol. Ser. 7: 18 (Nevada Test Site). —Wheeler and Wheeler, 1973.
Ants of Deep Canyon, pp. 101-102.

Iridomyrmex pruinosus
***authority mismatch
pruinosus (Roger). N. Y. to Fla., w. to Wis., Ohio, Tex., N. Mex.; W. Indies.

Ecology: Nests are in exposed soil with craterlike mounds or under the cover of objects.
A house pest in the Gulf Coast states.
Tapinoma pruinosum Roger, 1863. Berlin Ent. Ztschr. 7: 165. ♂.

Taxonomy: Smith, 1929. Jour. Econ. Ent. 22: 241. —Cole, 1940. Amer. Midland Nat. 24: 64-65.
—Cole, 1942. Amer. Midland Nat. 28: 372. — Santschi, 1930. Soc. Roy. Ent. d'Egypte, Bui.
23: 81. —Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 185. (larva). —Wheeler
and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 729 Oarva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 389. —Smith, 1930. Fla Ent. 14: 5.
— Wheeler, 1911. Harvard Univ., Mus. Comp. Zool., Bui. 54: 497. — Warter, Moser, and
Blum, 1962. La. Acad. Sci., Proc. 25: 42^6 (correlation between night hawk feeding and
nuptial flights of pruinosus pruinosus). — Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:
54-55 (economic importance).

Morphology: Blvun, Warter, Monroe, and Chidchester, 1963. Jour. Ins. Physiol. 9: 881-885
(chemical releasers of social behavior - methyl-n-amyl ketone).

Genus FORELIUS Emery

Forelius Emery, 1888. Ztschr. f. Wiss. Zool. 46: 389.

Type-species: Iridomyrmex mccooki Forel. Monotypic.

A small New World genus.

Taxonomy: Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 729 Oarvae).

Forelius foetidus (Buckley)
Ark., Kans., Okla., Tex., Colo. w. to Calif.; Mexico. Ecology: Nests are
under objects or in exposed soil with small craters of soil Incorrectly reported from D. C.
Formica foetida Buckley, 1866. Ent. Soc. Phila., Proc. 6: 167. ♂, ♀.
Iridomyrmex McCooki Forel, 1878. Soc. Vaud. des Sci. Nat., Bui. 15: 382.
Forelius MacCooki race Andrei Forel, 1912. Soc. Ent. de Belg., Mem. 20: 44. ♂.

Taxonomy: Forel, 1886. Soc. Ent. de Belg., Bui. (C. R.) 30: 39. —Wheeler and Wheeler, 1974
(1973). Pan-Pacific Ent. 49: 401 (larva).

Biology: Cole, 1937. Ent. News 48: 137. —Mallis, 1941. South. Calif. Acad. Sci., Bui. 48: 137.
— Lindquist, 1942. Jour. Econ. Ent. 35: 850-852. —Gregg, 1963. Ants of Colo., pp. 434-436.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 102-103.

Genus CONOMYRMA Forel

Conomyrma Forel, 1913. Rev. Zool. Africaine 2: 350.

Type-species: Prenolepis pyramica Roger. Desig. by Santschi, 1922.




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Conomyrma subg. Biconomyrma Kusnezov, 1952. Acta Zool. Lilloana (Tucuman) 10:
429-430.

Type-species: Dorymyrex pyramicus var. brunnea Forel. Desig. by Kusnezov, 1959.

A New World genus. The Nearctic forms construct nests in the soil which are surmounted by
Irregular or craterlike mounds of soil. They are usually found in op)en sunny areas in situations
where other ant species find conditions intolerable. The workers are predaceous, active and
agressive and emit a fluid with a disagreeable odor. The species below have previously been as-
signed to the genus Dorymyrmex Mayr, a genus now restricted to South America. The taxa of
this genus occurring in the Gulf coast states are not yet clearly imderstood.

Taxonomy: Gallardo, 1916. Buenos Aires Mus. Nac. de Hist. Nat., An. 28: 1-130. — Santschi,
1922. Soc. Vaud. des Sci. Nat., Bui. 54: 365. —Kusnezov, 1952. Acta Zool. Lilloana
(Tucuman) 10: 427, 429, 433^38. —Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53:
183. —Kusnezov, 1959. Zool. Anz. 162: 38-51. — Snelling, 1973. Los Angeles Co. Mus.,
Contrib. Sci. 238: 1-6 (in the U. S.).

Conomyrma bicolor (Wheeler)
Tex., N. Mex., Utah, Ariz., Nev., Calif.; Mexico. Ecology: Nests are in
exposed situations in the soil and are usually surmounted by irregular or craterlike
mounds.
Dorymyrmex pyramicus var. bicolor Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 342.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 372. —Cole, 1957. N. Y. Ent. Soc., Jour. 65:
130. —Wilson, 1957. Psyche 64: 76.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 76. — Eckert and Mallis, 1937. Calif.

Agr. Expt. Sta. Cir. 342: 19. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 18

(Nevada Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 103-104.

Conomyrma flavopecta (Smith)
Fla. Occurrence in other Gulf Coast states is not certain.

Dorymyrmex pyramiciLS Jlavopectus Smith, 1944. Fla. Ent. 27: 15. 2 .

Conomyrma insana (Buckley)
N. C. to Fla., w. to 111., N. Dak., Oreg., Calif.; Mexico; W. Indies. Ecology:

Nests are in soil in open sunny places and are surmoimted by an irregular or craterlike
mound. A pest species mainly in the South where it commonly enters houses and builds
ugly mounds on lawns. Pyramid ant. Records in the literature for Dorymyrmex
pyramicus (Roger) in N. Amer. should refer to this species; Conomyrma pyramica
(Roger) is a valid species but is found only in S. Amer.

Formica insana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 165. $ , 9 .

Dorymyrmex flavus McCook, 1879. In Comstock, Rpt. on Cotton Insects, p. 186. ♀.

Dorymyrmex pyramicus var. nigra Pergande, 1895. Calif. Acad. Sci., Proc. 5: 871. ♂.

Dorymyrmex pyramica brunnea var. antillana Forel, 1911. Deut. Ent. Ztschr., p. 306.

Dorymyrmex pyramicus var. smithi Cole, 1936. Ent. News 47: 120. ♂.

Conomyrma {'? Biconomyrma) wheeleri Kusnezov, 1952. Acta Zool. Lilloana (Tucuman) 10:
438. ♂.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System 8: 331-332 (male). —Cole, 1940. Amer.
Midland Nat. 24: 61. —Cole, 1942. Amer. Midland Nat. 28: 371. —Smith, 1936. Puerto Rico
Univ., Jour. Agr. 20: 861, 864. — Buren, 1944. Iowa State Col., Jour. Sci. 18: 291. —Wheeler
and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 183 (larva). —Wilson, 1967. Psyche 64: 76
(sympatry of bicolor and pyramica). — Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci.
238: 5 (syn.).

Biology: Wheeler, 1910. Ants, pp. 146, 210, 205, 426. —Smith, 1924. Ent. News 35: 82.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31: 293, 305. —Eckert and MaUis, 1937. Calif. Agr.
Expt. Sta. Cir. 342: 19. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 155-157.
—Gregg, 1963. Ants of Colo., pp. 432-434. —Wilson, 1967. Psyche 64: 76. —Smith, 1965. U.
S. Dept. Agr., Tech. Bui. 1326: 56-57 (economic importance). —Cole, 1966. Brigham Young
Univ., Sci. Bui., Biol. Ser. 7: 18 (Nevada Test Site). —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 105-106.

Morphology: Metcalf and Metcalf, 1970. Ent. Soc. Amer., Ann. 63: 34-35 (effects of isomeres
of 2-heptanone on alarm behavior). —Blum and Warter, 1966. Ent. Soc. Amer., Ann. 59:
774-779 (alarm and digging behavior; isolation of 2-heptanone).




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Genus TAPINOMA Foerster

Tapinoma Foerster, 1850. Hym. Stud. 1: 43.

Type-species: Tapinoma colliyia Foerster. Monotypic.
Micromyrma Dufour, 1857. Soc. Ent. de France, Aim. 5: 60.

Type-species: Micromyrma pygmaea Dufour. Monotypic.
Tapinoma subg. Tapinoptera Santschi, 1925. Eos 1: 348.

Type-species: Tapinoma vexatum Santschi. Monotypic.

At least one of the three North American species is introduced. Nests are found in a variety
of situations and workers commonly attend honeydew-excreting insects. All of the species emit a
substance with an odor similar to that of butyric acid.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 172 (larvae).

Morphology: Torossian, 1%0. Insectes Sociaux 7: 171-174 (physiology of the Palearctic

Tapinoma erraticum (Latreille)). —Weber, 1961. Ent. Soc. Wash., Proc. 63: 217-218 (use of
poison).

Tapinoma litorale Wheeler
S. Fla.; W. Indies. Ecology: Apparently arboreal, nesting in twigs of trees
and bushes, hollow culms, or between leaves.
Tapinoma litorale Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 109. 5, 9, 6.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 861-862.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 150. —Wheeler, 1932. N. Y. Ent. Soc.,

Jour. 40: 13.

Tapinoma melanocephalum (Fabricius)
Fla., where it is established; occasionally found in greenhouses
or heated buildings in other parts of the continent such as Ont., Iowa., N. Mex., CaUf.;
tropicopolitan. Ecology: Highly adaptable in nesting habits; found in soil, rotten wood,
decayed parts of trees, under bark, in plant cavities, houses, and greenhouses. Workers
are fond of honeydew but also feed on live and dead insects. A house-infesting ant.
Introduced; widely distributed in the tropical regions of the world by commerce.

Formica melanocephala Fabricius, 1793. Ent. System. 2: 353. ♂.

Formica nanfl Jerdon, 1851. Madras Jour. Lit. Sci. 17: 125. ♂.

Myrmica pelludda Smith, 1857. Linn. Soc. London, Jour., Zool. 2: 71. ♂.

Formica familiaris Smith, 1860. Linn. Soc. London, Jour., Zool., Sup. 4: 96. ♂.

Tapinoma {Micromyrma) melanocephalum war. australis Santschi, 1928. Insects of Samoa
5:53. ♂.

Taxonomy: Smith, 1928. Ent. Soc. Amer., Ann. 21: 311. —Smith, 1936. Puerto Rico Univ.,
Jour. Agr. 20: 861-862. —Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 197
(larva). — WUson and Taylor, 1967. Pacific Ins. Monog. 14: 79-82 (Polynesia).

Biology: Wheeler, 1910. Ants, pp. 154, 156. — Marlatt, 1928. U. S. Dept. Agr. Farmers' Bui.
740: 6. —Phillips, 1934. Hawaii Univ., Expt. Sta. Pineapple Prod. Coop. Assoc., Bui. 15:
20-21. —King, 1948. Iowa Acad. Sci., Proc. 55: 395. — Kempf, 1961. Studia Ent. 4: 520.
—Brown, 1964. Ent. News 75: 14-15. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326:
59-60 (economic importance). — Tamsitt and Fox, 1966. Ent. Soc. Wash., Proc. 68: 268
(attacking lab. flea colony). —Gomez-Nunez, 1971. Jour. Med. Ent. 8: 735-737 (an inhibitor
of Rhodimis prolixus populations).

Morphology: Weber, 1961. Ent. Soc. Wash., Proc. 63: 218 (employment of venom).

Tapinoma sessile (Say)
N. S., Que. s. to Fla., w. to Wash., Calif.; Mexico. Ecology: A very adaptable ant
found in a wide variety of habitats. Most nests are in the soil beneath objects but also
under bark, in stumps, plant cavities, insect galls, refuse pUes, and bird and mammal
nests. Colonies may contain thousands of individuals and numerous reproductive
females. A common and troublesome house-infesting ant. Food consists mainly of
honeydew and workers attend honeydew-excreting insects. Odorous house ant.

Formica sessilis Say, 1836. Boston Jour. Nat. Hist. 1: 287. ♂, ♀.

Tapinoma bareale Roger, 1863. Berlin. Ent. Ztschr. 7: 165. ♂, ♀.

Formica gracilis Buckley, 1866. Ent. Soc. Phila., Proc. 6: 158. 9, 9.

Formica parva Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. ♀.




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Tapinoma bareale Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 238. 9,9.
Preocc. by Roger, 1863.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 196 (larva). — Francoeur
and Beique, 1966. Canad. Ent. 98: 143 (Provancher types).

Biology: Smith, 1928. Ent. Soc. Amer., Ann. 21: 307-329. — Metcalf and Flint, 1939.
Destructive and Useful Insects, Ed. 2: 770. —Weber, 1941. Canad. Ent. 73: 140-141.
— Kannowski, 1959. Insectes Sociaux 6: 126-129. —Wheeler and Wheeler, 1963. Ants of N.
Dak., pp. 157-160. —Gregg, 1963. Ants of Colo., pp. 443-447. ^Judd, 1964. Canad. Ent. 96:
990 (in galls on goldenrod). —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326: 57-59
(economic importance). — Bobb, 1965. Jour. Econ. Ent. 58: 925 (as predator of Neodiprion
pratti pratti (Dyar)). — Kulman, 1965. Jour. Econ. Ent. 58: 70 (as predator of Malacosoma
americanum (F.)). —Wang and Brook, 1970. Jour. Econ. Ent. 63: 1971-1973 (toxicological
and biological studies). — Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 106-107.

Morphology: Crozier, 1970. Canad. Jour. Genet, and Cytol. 12: 541-546 (pericentric
rearrangement polymorphism).

Unplaced Taxa of Douchoderinae

Bothriomyrmex dimmocki Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 417. 5,9. Mass. (Mt.

Tom, Springfield).
Formica masonia Buckley, 1866. Ent. Soc. Phila., Proc. 6: 165. 5 . Tex. (Fort Mason).
Formica (Hypochira (.')) subspinosa Buckley, 1866. Ent. Soc. Phila., Proc. 6: 169. $. Tex.

Subfamily FORMICINAE

This subfamily is the predominant group of ants in North America, most common in the north
and in the mountains with their numbers decreasing rapidly toward the southern part of the
United States. They are recognized by the single segmented pedicel, lack of a constriction
between the first and second gastric segments, and the round acidopore, usually with a circle of
hairs.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 126-171, 175-217 (larvae).
—Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ami. 61: 205-222 (larvae). —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ami. 63: 648-656 (larvae).

Biology: Kannowski, 1963. Symp. (Jenet. et BioL Ital. 12: 74-102 (flight activities). —Wilson
and Regnier, 1971. Amer. Nat. 105: 279-289 (evolution of alarm-defense system).

Morphology: Blum and Wilson, 1964. Psyche 71: 28-31 (anatomical source of trail substances
in formicine ants). — Hung, 1969. Ent. Soc. Amer., Ann. 62: 455-456 (chromosome numbers
of various spp.).

Tribe PLAGIOLEPIDINI

Genus PLAGIOLEPIS Mayr

Plagiolepis Mayr, 1861. Eurp. Formicid., p. 42.

Type-species: Formica pygmaea Latreille. Monotypic.
Plagiolepis subg. Anacantholepis Santschi, 1914. Meddel. Groteborgs Mus. ZooL 3: 36.

Type-species: Plagiolepis (Anacantholepis) decora Santschi. Monotypic.
Aporxymyrmex Faber, 1969. Pflanzenschutzber. 39: 52.

Type-species: Aporomyrmex am.peloni Faber. Orig. desig.
Plagiolepis subg. Paraplagiolepis Faber, 1969. Pflanzenschutzber. 39: 65.

Type-species: Plagiolepis xene Starcke. Monotypic.

An Old World genus with a single introduced species found in the United States.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 129, 132 (larvae). — Brown.
1973. In Meggers, et al.. Tropical forest ecosystems in Afr. and S. Amer., pp. 178-185.




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Plagiolepis alluaudi Emery
Calif. (Catalina Is.); W. Indies, Pacific Islands, apparently pantropical.

Introduced, probably native to Africa from where it has been widely distributed by

commerce.
Plagiolepis alltmitdi Emery, 1894. Soc. Ent. de France, Ann. 63: 71. ♂.
Plagiolepis mactavishi Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 166. § .
Plagiolepis foreli Santschi, 1920. Soc. Vaud. des Sci. Nat., Bui. 53: 165. ♂.
Plagiolepis foreli var. omata Santschi, 1920. Soc. Vaud. des Sci. Nat., Bui. 53: 166. ♂.
Plagiolepis foreli Mann, 1921. Harvard Univ., Mus. Comp. ZooL, Bui. 64: 473. ♂. Preocc. by

Santschi, 1920.
Plagiolepis augusti Emery, 1921. Soc. Ent. de Belg., Ann. 61: 317. N. name for foreli

Mann.

Taxonomy: Smith, 1957. N. Y. Ent. Soc., Jour. 65: 196 (also biol. notes). —Brown, 1958. Acta
Hym. 1: 49. —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 85-86 (Polynesia).

Biology: Phillips, 1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc., Bui. 15: 17-18
(in Hawaii). —Taylor and Wilson, 1961. Psyche 68: 143.

Genus ACROPYGA Roger

Acropyga Roger, 1862. Berlin. Ent. Ztschr. 6: 242.

Type-species: Acropyga acutiventris Roger. Monotypic.
Acropyga subg. Rhizomyrma Forel, 1893. Ent. Soc. London, Trans., p. 347.

Type-species: Acropyga (Rhizomyrma) goldii Forel. Desig. by Wheeler, 191 1.
Acropyga subg. Atopodon Forel, 1912. Rev. Suisse de Zool. 20: 771.

Type-species: Acropyga {Atopodon) inerzae Forel. Desig. by Wheeler, 1913.
Acropyga subg. Malacomyrma Emery, 1922. Mus. Nat. Hung., Ann. 19: 109.

Type-sp>ecies: Acropyga silvestrii Emery. Monotypic.

The small yeUow ants of this genus are subterranean in habit and are fond of honeydew. Ac-
cording to Weber (1944) all the New World species may be obligate coccidophiles. Some species
may be of economic importance because they tend and disperse coccids on the roots of coffee
plants.

Revision: Weber, 1944. Ent. Soc. Amer., Ann. 37: 89-122 (New World spp.; also biol. and
association with coccids).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 136 Oarvae). — Snelling,
1973. Los Angeles Co. Mus., Contrib. Sci. 236: 6-7 (first record of genus in U. S.).

Acropyga epedana Snelling
Ariz. (Montezuma Pass, 5800 ft., Huachuca Mts., Cochise Co.) Ecology:
Specimens were found imder a stone.
Acropyga epedana Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 236: 7-8. ♀.

Tribe MYRMELACHISTINI

Genus MYRMELACHISTA Roger

Mynnelachista Roger, 1863. Berlin. Ent. Ztschr. 7: 162.

Type-species: Myrmelachista kraatzii Roger. Monotypic.
Myrmelachista subg. Decamera Roger, 1863. Berlin. Ent. Ztschr. 7: 166. Preocc. by
Mulsant, 1842.

Type-species: MyrmeUichista (Decamera) nigella Roger. Monotypic.
Aphomomyrmex subg. Neaphomus Menozzi, 1935. Zool. Jahrb., Abt. f. System 67: 324.
Synonymy questionable.

Type-Sf>ecies: Aphomomyrmex (Neaphxytmis) goetschi Menozzi. Monotypic.
Hincksidris Donisthorpe, 1944. Ent. Monthly Mag. 80: 59. N. name for Decamera Roger.

A Neotropical genus represented in the United States by a single species probably introduced
into Florida.

Taxonomy: Wheeler, 1934. Harvard Univ., Mus. Comp. Zool., Bui. 77: 187-206. —Wheeler and
Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 129, 137 Oarvae). — Kusnezov, 1959. Acta ZooL
Lilloana (Tucuman) 17: 379.




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Myrmelachista ramulorum Wheeler
Fla. (Highland City, "Polk Co.); W. Indies. Ecology: Specimens in Florida
collected on sweet orange. Probably introduced.
Myrmelachista ambigua ramulorum Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:

155.5,9, <5.

Taxonomy: Wheeler, 1934. Harvard Univ., Mus. Comp. Zool., Bui. 77: 189. —Smith, 1936.
Puerto Rico Univ., Jour. Agr. 20: 872-873 (also biol notes). —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 47: 138 (larva).

Biology: Anonymous, 1968. FAO Plant Protect. Bui. 16: 53 (Fla. record; biol).

Tribe BRACHYMYRMECINI
Genus BRACHYMYRMEX Mayr

Brachymyrmex Mayr, 1868. Soc. Nat. Modena, Ann. 3: 163.

Type-species: Brachymyrmex patagonicus Mayr. Monotypic.
Brachymyrmex subg. Bryscha Santschi, 1925. Buenos Aires Mus. Nac. de Hist. Nat., An.
31: 652.

Type-species: Brachymyrmex pilipes Mayr. Orig. desig.

Most species of this New World genus are found south of the United States. They usually
form small colonies in the soil under various objects and are easily transported by commerce in
soil or plants. They are fond of honeydew and attend honeydew excreting insects.

Revision: Wheeler, 1903. Psyche 10: 102-103. —Santschi, 1923. Buenos Aires Mus. Nac. de
Hist. Nat., An. 31: 650-674.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 129, 139 (larvae).

—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 652 (larvae, Braehymyrmecini).

Biology: Amaud and Quate, 1951. Pan-Pacific Ent. 27: 171 (note on swarming).

Brachymyrmex depilis Emery
N. S. s. to Fla. w. to B. C, Calif. Ecology: Nests are small and inconspicuous
and are in the soil under stones, rotten wood, or other objects.
Brachymyrmex heerii depilis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 635. ♂.
Brcu:hym,yrm£X nanellus Wheeler, 1903. Psyche 10: 102. 5, Brachym,yrm^x depilis flavescens Grtmdmann, 1952. Kans. Ent. Soc., Jour. 25: 117. ♂.

Taxonomy: Santschi, 1923. Buenos Aires Mus. Nac. de Hist. Nat., An. 31: 664. —Cole, 1940.
. Amer. Midland Nat. 24: 65. —Gregg, 1944. Ent. Soc. Amer., Ann. 37: 469. —Cole, 1953.
Ent. News 64: 266. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 139 (larva).
—Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 210 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 389. —Smith, 1927. Ent. News 38:
313. —Dennis, 1938. Ent. Soc. Amer., Ann. 37: 469. — Headley, 1952. Ent. Soc. Amer., Ann.
45: 436, 439. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 161-163. —Gregg, 1963.
Ants of Colo., pp. 447^449. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp.
108-109.

Brachymyrmex obscurior Forel
Fla., s. Tex.; Mexico, Central Amer., W. Indies. Possibly introduced into U. S.
Brachymyrmex heeri var. obscurior Forel, 1893. Ent. Soc. London, Trans., p. 345. 5, 2, 6.

Taxonomy: Santschi, 1923. Buenos Aires Mus. Nac. de Hist. Nat, An. 31: 654, 666. —Smith,
1936. Puerto Rico Univ., Jour. Agr. 20: 866-867 (also biol. notes). —Brown, 1957. Harvard
Univ., Mus. Comp. Zool., Bui. 116: 237.

Biology: Plank and Smith, 1940. Puerto Rico Univ., Jour. Agr. 24: 60.

Tribe CAMPONOTINI
Genus CAMPONOTUS Mayr

This large genus of ants is found throughout North America and most other parts of the
world. They are commonly referred to as carpenter ants, especially members of the subgenus
Camponotus, because of their habit of nesting in wood. However, many species nest in the soil,




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under stones, or in hollow twigs or branches.

Many subgeneric groupings have been proposed for Camponotus on a worldwide basis, many
of which may or may not be valid or may eventually represent distinct genera. Brown (1973)
gave 54 genus-group names to be considered with Camponotiis. Seven subgenera have been
recognized in North America, and these are separated here as in the past literature.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 667-682. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 295-354.

Taxonomy: Emery, 1920. Rev. Zool. Afr. 8: 16-19 (subgenera). —Wheeler, 1921. Psyche 28:
17. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 181 (larvae). —Wheeler and
Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 216 (larvae). —Wheeler and Wheeler 1970. Ent.
Soc. Amer., Ann. 63: 649 (larvae). — Brown, 1973. In Meggers, et al., Tropical forest
ecosystems in Afr. and S. Amer., pp. 178-185 (genus-group names).

Biology: Smith, 1942. Ent. News 53: 133-135 (effect of reduced food supply on stature).
—Smith, 1942. Tenn. Acad. Sci., Jour. 17: 367-373 (polymorphism). — Riordan, 1960.
Insectes Sociaux 7: 353-355 (location of nests by a radioactive isotope). — Sanders, 1964.
Canad. Ent. 96: 894-909 (biol. of carpenter ants in N. B.). —Patrick, 1969. Amer. Midland
Nat. 82: 605-610 (changes in carpenter ants harboring dicrocoeliid metacercariae).
— Sanders and Baldwin, 1969. Canad. Ent. 101: 416-418 (Iridiiun-192 as a tag for carpenter
ants). — Sanders, 1970. Ecology 51: 865-873 (distribution of carpenter ant colonies in
spruce fir forests in Ont.). — Ayre and Blum, 1971. Physiological Zool. 44: 77-83 (attraction
and alarm). —Sanders, 1972. Canad. Ent. 104: 1681-1687 (seasonal and daily activity
patterns in n.w. Ont.). —Sanders, 1973. Ent. Soc. Ontario, Proc. 102: 13-16 (aggregation of
alate carpenter ants in Ont.).

Genus CAMPONOTUS Subgenus CAMPONOTUS Mayr

Camponottts Mayr, 1861. Die Europaischen Formiciden, pp. 25, 35.

Type-species: Formica ligniperda Latreille. Desig. by Bingham, 1903.

Most species of this subgenus build their nests in decaying wood. Some are common house in-
festing ants that make their nest in the timber and woodwork of buildings and tunnel nest
passages into adjacent wood. Dry, sound wood is rarely attacked, and the presence of nests of
carpenter ants in buildings is usually evidence that the wood was damp and partially decayed.

Camponotus americanus Mayr
Ont. s. to Fla. w. to Mich., Iowa, Mo., Okla., Tex. Ecology: Prefers to nest in
the soil, usually under stones or rotten logs.
CaTnponottts amencamis Mayr, 1862. Zool.-Bot. (Jesell. Wien, Verh. 12: 661. ♂, ♀.
Camponotus (Cam,ponotus) castaneus stirps rufinasis Santschi, 1936. Rev. d. Ent. 6: 204.

5.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 323-325 (each caste). —Cole, 1940.
Amer. Midland Nat. 24: 82, 84 (also biol. notes). —Gregg, 1944. Ent. Soc. Amer., Ann. 37:
477. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 185 Garva). —Wheeler and
Wheeler, 1958. Ent. Soc. Amer., Ann. 61: 216 Oarva).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 273, 275, 300-301.

Morphology: Ayre and Blum, 1971. Physiological Zool. 44: 77-83 (attraction and alarm by
pheromones).

Camponotus ferrugineus (Fabricius)
Mass., N. Y. s. to Ga. w. to Mich., III., Nebr., Kans. Ecology: Nests are
located in and beneath well-rotted logs and stumps with galleries often extending into
the soil. They have also been found in dead standing trees and occasionally in moist or
faulty wood in buildings. Red carpenter ant.
Formica ferruginea Fabricius, 1798. Sup. Ent. System., p. 279. ♂, ♀.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 668. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 299, 338-339 (each caste; biol. notes). —Wheeler, 1917. Conn. State Geol. and
Nat. Hist. Survey Bui. 22: 600-601. —Cole, 1940. Amer. Midland Nat. 24: 86. —Brown,
1950. Ent. News 61: 158-160. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 188
(larva).




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Biology: Pricer, 1908. Biol. Bui. 14: 177-218. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:
301-302. —Williams, 1961. Ohio Jour. Sci. 61: 279 (habits; nest). —Smith, 1965. U. S. Dept.
Agr., Tech. Bui. 1326, pp. 67-69 (economic importance).

Morphology: Tanquary, 1913. 111. State Lab. Nat. Hist., Bui. 9: 454-475 (embryology).

Camponotus herculeanus (Linnaeus)
Newfoundland (Labrador and insular) w. to Alaska s. to N. Y., Pa.,
Wis., Minn., N. Dak., Colo., N. Mex., Utah, Oreg.; Eurasia. Ecology: Probably the
dominant ant in the forests of boreal and alpine N. Amer. Large colonies are found in
rotting logs and stumps, especially conifers.
Formica herculeana Linnaeus, 1758. Syst. Nat., Ed. 10, p. 579. ♂.

Camponotus herculeanus var. WhympeH Forel, 1902. Ent. Soc. London, Trans., p. 699. ♂.
9.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 330-333 (each caste; biol. notes).
—Cole, 1942. Amer. Midland Nat. 28: 387. — Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bui. 104: 367. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 185-187
(larva). — Yasumatsu and Brov*m, 1957. Kyushu Univ., Fac. Agr., Jour. 11: 45. — Amoldi,
1967. Zool. Zhur. 46: 1817-1818. —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63:
651 (larva).

Biology: Jones, 1929. Colo. Agr. Expt. Sta. Bui. 341, 96 pp. (tends 19 spp. in 6 gen. of aphids
on 12 spp. of host plants). —Gregg, 1946. Amer. Midland Nat. 35: 753. —Brown, 1949. Ent.
News 60: 99. — Holldobler, 1950. Ztschr. f. Angew. Ent. 31: 583 (biol. and habits in
Europe). —Weber, 1950. Amer. Ent. Soc., Trans. 76: 188. — Perttunen, 1955. Ann. Ent.
Fenn. 21: 38 (reactions to air humidity). —Brown, 1955. Ent. News 66: 47-48. — HoUdobler,
1961. Insectes Sociaux 8: 14 (rhythmical behavior). —Holldobler, 1962. Waldhygiene 4: 228
(nest types, ecomomic importance in Germany). — Holldobler, 1962. Ztschr. f. Angew. Ent.
49: 338. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 164-165. —Gregg, 1963. Ants of
Colo., pp. 656-660. —Sanders, 1964. Canad. Ent. 96: 894 (biol. and habits, N. B.). —Sanders,
1972. Canad. Ent. 101: 1618-1687 (seasonal and daily activity patterns, Ont.).

Morphology: Ayre, 1963. Ent. Expt. and Appl. 6: 165-170 (feeding behavior and digestion).
— Ayre and Blum, 1971. Physiological Zool. 44: 77-83 (attraction and alarm by
pheromones).
laevig^atus (Smith). Mont., Colo., N. Mex. w. to B. C, Oreg., Calif.; Mexico. Ecology: Foimd in
wooded and forested areas where it nests in rotting logs and stumps. Occasionally foimd
in buildings.
Formica laevigata Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 55. ♂, ♀.

Taxonomy: Wheeler, 1910. N. Y. Acad. ScL, Ann. 20: 299, 327-330 (each caste; biol. notes).
—Cole, 1942. Amer. Midland Nat. 28: 387-388. —Wheeler and Wheeler, 1968. Ent. Soc.
Amer., Ann. 61: 216 Garva).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 556. — Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342" 29. — Fumiss, 1944. Oreg. Agr. Expt. Sta. Cir. 158: 1-12.
—Gregg, 1963. Ants of Colo., pp. 660-663.

Camponotus modoc Wheeler
S. Dak., Colo. w. to B. C, Oreg., Calif. Ecology: Found in forested areas

where it makes its nests in rotting logs and stumps. Has been considered a subspecies
of herculeanus or pennsylvanicus by various authors.
Camponotus (Camponotus) herculeanus var. modoc Wheeler, 1910. N. Y. Acad. Sci., Ann.
20:299,333. 5, 9, 6.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 387. —Brown, 1950. Ent. News 61: 158.

Biology: Wheeler, 1917. Amer. Acad. Arts and ScL, Proc. 52: 557. —Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 29. —Mallis, 1941. South. Calif. Acad. Sci., BuL 40: 90.
—Fumiss, 1944. Oreg. Agr. Expt. Sta. Cir. 158: 1-12. -Gregg, 1963. Ants of Colo., pp.
665-669.

Camponotus novaeboracensis (Fitch)
N. S., Que. s. to Va. w. to B. C, Oreg., Utah, Colo. Ecology: Prefers
wooded areas where it normally nests in rotting logs and stumps. Sometimes a house
pest.
Formica Novaeboracensis Fitch, 1855. N. Y. State Agr. Soc., Trans. 14: 766. ♀.




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CampoTwtus herculeanus ligniperdus var. pictus Forel, 1879. Soc. Vaud. des Sci. Nat., Bui.

16:59. 9, 9, 6.
Camponotus herculeanus ligniperdus var. noveboracensis Forel, 1899. Soc. Ent. de Belg.,

Ann. 43: 447. Emend.
Camponotus herculeanus ligniperda var. rubens Wheeler, 1906. Psyche 13: 41. ♀, ♂.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 340 (each caste). — Buren, 1944.
Iowa State Col., Jour. Sci. 18: 293. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46:
182-186 (larva). —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larva).

Biology: Jones, 1929. Colo. Agr. Expt. Sta. Bui. 341: 96 pp. (attends aphids). —Gregg, 1944.
Ent. Soc. Amer., Ann. 37: 457-458. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11:
250-251. — Kannowski, 1959. Insectes Sociaux 6: 134-135. —Wheeler and Wheeler, 1963.
Ants of N. Dak., pp. 165-168. —Gregg, 1963. Ants of Colo., pp. 663-665. —Sanders, 1964.
Canad. Ent. 96: 896 (biology and habits, N. B.). — Gotwald, 1968. N. Y. Ent. Soc., Jour. 76:
278-296 (food gathering behavior). —Sanders, 1972. Canad. Ent. 104: 1681-1687 (seasonal
and daily activity pattern, Ont.).

Morphology: Smith, 1942. Tenn. Acad. Sci., Jour. 17: 368.

Camponotus pennsylvanicus (DeGeer)
N. B., Que. s. to Fla. vi. to N. Dak., Tex. Ecology: Nests are found in
live and dead trees, rotting logs and stumps, and in wood products such as fences,
telegraph poles, and buildings. A common and important house infesting ant because of
its adaptability to nest in woodwork of buildings. It commonly feeds on household foods,
and it is annoying. Nests are started in buildings usually in moist or faulty wood. The
first native North American ant to be described. Black carpenter ant.

Formica pensylvanica (!) DeCJeer, 1773. Mem. Serv. Hist. Ins., v. 3, p. 603. §, 9, 6.

Camponotus herculeanus herculeanus herculeano-pennsylvanicus Forel, 1879. Soc. Vaud.
des Sci. Nat., Bui. 16: 57. 5 (?).

Camponotus herculeanus pennsylvanicus var. mahican Wheeler, 1910. N. Y. Acad. Sci.,
Ann. 20: 299, 338. N. name for herculeano-pennsylvanicus Forel.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 335-336 (each caste). —Wheeler,
1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22: 600. —Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 293. — Townsend, 1945. Ky. Agr. Expt. Sta. Cir. 59: 1-27 (references to 1945).
—Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 187-188 (larva). —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Arm. 63: 651 flarva).

Biology: McCook, 1877. Amer. Ent. Soc., Trans. 6: 253-296. — Pricer, 1908. Biol. Bui. 14:
177-218. — Herrick, 1914. Insects Injurious to the Household and Annoying to Man, pp.
177-178. —Gibson, 1916. Canad. Ent. 48: 365-366. —Graham, 1918. Minn. State Ent. Rpt.
17: 32-40 (as destroyers of sound wood). —Back, 1937. U. S. Dept. Agr. Leaflet 147. —Van
Pelt, 1958. Tenn. Acad. Sci., Jour. 33: 120-122 (parasitism by a fungus). —Wheeler and
Wheeler, 1963. Ants of N. Dak, pp. 168-170. —Sanders, 1964. Canad. Ent. 96: 899 (biology
and habits, N. B.). —Smith, 1965. U. S. Dept. Agr. Tech. Bui. 1326, pp. 63-67 (economic
importance). —Sanders, 1972. Canad. Ent. 104: 1681-1687 (seasonal and daily activity
patterns, Ont.).

Morphology: McCook, 1878. Phila Acad. Nat. Sci., Proc. 30: 15-19. — Fielde, 1903. Biol. Bui. 5:
320-325. —Fielde and Parker, 1904. Phila. Acad. Nat. Sci., Proc. 56: 642-649. —Forbes,
1938. Ent. Soc. Amer., Ann. 31: 181-195 (anatomy and histology of worker). —Smith, 1942.
Tenn. Acad. Sci., Jour. 17: 367-373. —Forbes, 1952. N. Y. Ent. Soc., Jour. 60: 157-171 (male
genitalia). —Forbes, 1954. Jour. Morph. 95: 523-548 (male reproductive system). —Forbes,
1956. Insectes Sociaux 3: 505-511 (male digestive tract). —Forbes and McFarlane, 1961. N.
Y. Ent. Soc., Jour. 69: 92-103 (comparative anatomy of digestive glands of female and
male). — Keister, 1963. Ent. Soc. Amer., Ann. 56: 33&-340 (trachael system). —Hermann
and Blum, 1968. Psyche 75: 216-227 (poison apparatus). — Ayre and Blum, 1971.
Physiological Zool. 44: 77-83 (attraction and alarm by pheromones).

Camponotus quercicola Smith
Calif. (Los Angeles Co. to San Diego Co.). Ecology: Nests in dead limbs of
Qu^ercvs agrifolia; nocturnal.
Camponotus (Camponotus) quercicola Smith, 1953. N. Y. Ent. Soc., Jour. 61: 211-214. ♀.




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Camponotus schaefferi Wheeler
S. Ariz. Ecology: Nests in dead oak limbs at elevations of about 5000 to
8000 feet.
Camponotus schaefferi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 88. 5 , 2 .

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 344-345 (also biol. notes).

Biology: Wheeler, 1917. Amer. Acad. Arts, and Sci., Proc. 52: 557.

Camponotus texanus Wheeler
Tex. Ecology: Nests have been found in oak logs.
Camponotus texanus Wheeler, 1903. Psyche 10: 108. 5 , 9 , 6.

Taxonomy: Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 90. —Wheeler, 1910. N. Y. Acad. Sci.,
Ann. 20: 300, 344 (also biol. note).

Genus CAMPONOTUS Subgenus TANAEMYRMEX Ashmead

Tanaeniyrmex Ashmead, 1905. Canad. Ent. 37: 384.

Type-species: Formica longipes (Jerstaecker. Grig, desig.
Myrmoturba Forel, 1912. Soc. Ent. de Belg., Mem. 20: 91.

Type-species: Formica m,aculata Fabricius. Desig. by Wheeler, 1913.

Most species of this subgenus nest in the soil under stones or other objects; occasionally nests
may be surmounted by a smaU crater. The ants rarely nest in wood, but, if so, the wood is
usually buried in the soil.

Camponotus acutirostris Wheeler
Tex., N. Mex., Ariz. Ecology: Nests have been found in the ground
under stones.
Cam,ponotus (Cam.ponotv^) acutirostris Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 298, 317.

Camponotus acutirostris var. clarigaster Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34:
420. ♂.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 561-562.

Camponotus castaneus (Latreille)
N. Y. s. to Fla. w. to Iowa, Okla., Tex. Ecology: Nests in rotting logs and
stumps, exposed soil, or in soil under objects. Occasionally enters buildings usually in
search of food.
Formica castanea Latreille, 1802. Hist. Nat. Fourmis, p. 118. 5, 9, cJ.
Formica m^llea Say, 1831. Descr. N. Spp. N. Amer. Ins. Found in La. by Jos. Barabino, p.

14. ♂.
Cam,ponotus clarus Mayr, 1862. Zool.-Bot. Gesell. Wien, Verb. 12: 660. § .

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 321-323. —Cole, 1940. Amer.
Midland Nat. 24: 82, 84.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 402. —Gregg, 1944. Ent. Soc. Amer.,
Ann. 37: 457, 477. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 60-62.

Camponotus dumetorum Wheeler
S. Calif. Ecology: Nests are common in the chaparral and are in the soU
usually surmounted by flat craters.
Camponotus (.Cam.ponotus) mnculatus dumetorum, Wheeler, 1910. N. Y. Acad. Sci., Ann.
20:354. ♀, ♂.

Taxonomy: Snelling, 1970. Ent. Soc. Wash., Proc. 72: 390-397.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 560. — MaUis, 1941. South. Calif.
Acad. Sci., Bui. 40: 90-91.

Camponotus festinatus (Buckley)
Tex., Colo., N. Mex., Ariz., Calif.; Mexico. Ecology: Nests in the ground

under stones, logs, or dried cow dung.
Formica festinata Buckley, 1866. Ent. Soc. Phila., Proc. 6: 165. ♂, ♀.
Camponotus {Cam.ponotus) fumidus var. pubicomis Emery, 1893. Zool. Jahrb., Abt. f.

System. 7: 668, 670. ♂.
Cam,ponotus fragilis Pergande, 1893. Calif. Acad. Sci., Proc. 4: 26. ♀.
Cam.ponotu3 (Camporwtus) fumidus var. spurcus Wheeler, 1910. N. Y. Acad. Sci., Ann. 20:

299,315. 9, 9.




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Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 312 (also biol. notes). — Snelling, 1968.
Ent. Soc. Wash., Proc. 70: 350-355. —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63:
651 (larva).

Biology: Wheeler, 1901. Amer. Nat. 35: 518, 520, 533.

Camponotus incensus Wheeler
Fla. (Pigeon Key near Miami).

Camponots (Tanaemyrmex) incensus Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 14. ♂.

Camponotus ocreatus Emery
N. Mex., Ariz., s. Nev., s. Calif.; Mexico. Ecology: Nests are found vmder
stones.
Camponotus (Camponotus) maculatus ocreatus Emery, 1893. Zool. Jahrb., Abt. f. System.

7: 668, 673. ♀.
Camponotus (Camponotus) acutirostris prvrnipilaris Wheeler, 1910. N. Y. Acad. Sci., Ann.
20:298,319. ♂, ♀.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 561-562. —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 661 flarva).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 272. —Cole, 1966. Brigham Young Univ., Sci.
Bui., Biol. Ser. 7: 20 (Nev. Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon,
p. 111.

Camponotus sansabeanus
***authority mismatch
bulimosus Wheeler. N. Mex., Ariz. Ecology: Nests in the ground under stones.
Camponotus (Camponotus) maculatus bulimosus Wheeler, 1910. N. Y. Acad. Sci., Ann. 20:
308. 5, 9, 6.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 560-561. —Cole, 1954. Tenn.
Acad. Sci., Jour. 29: 272.

Camponotus sansabeanus
***authority mismatch
sansabeanus (Buckley). Ark., La. w. to Colo., Utah, Ariz., s. Calif. Ecology: Nests
under rocks and stones in dry woods.
Formica San Sabeana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 167. 5, 9, 6.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 672-673. —Wheeler, 1910. N. Y.
Acad. Sci., Ann. 20: 298, 307-308 (also biol. notes). —Cole, 1942. Amer. Midland Nat. 28:
387. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 196 Oarva).

Biology: Wheeler, 1901. Amer. Nat. 35: 518, 520, 533. —Wheeler, 1910. Ants, pp. 349, 393.
—Gregg, 1963. Ants of Colo., pp. 669-671. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, p. Ill (half of collections belong to subsp. torrefactus).

Morphology: Wheeler, 1910. Ants, pp. 24, 49.

Camponotus sansabeanus
***authority mismatch
torrefactus Wheeler. Colo., Ariz., Utah, Nev. Ecology: Nests are found under
stones.
Camponotus (Cam.ponotus) maculatus sansabeanus var. torrefactus Wheeler, 1910. N. Y.
Acad. Sci., Ann. 20: 298, 308. 5, d.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 387, 388 (also bioL note).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 559.

Camponotus semitestaceus Emery
Okla., Tex. w. to Wash., Oreg., Calif.; Mexico. Ecology: Nests are foimd
under stones or in soil surmounted by low craters. Most U. S. records and references in
the literature to maccooki Forel refer to semitestaceus; maccooki is apparently
restricted to Guadalupe Is.
Camponotus (Camponotus) mxiculatus vicinus var. semitestaceus Emery, 1893. Zool.
Jahrb., Abt. f. System. 7: 668, 672. ♂.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 298, 304, 306. —Snelling, 1970. Ent. Soc.
Wash., Proc. 72: 390-397 (separation of maccooki and semitestaceus).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 345. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 559. — Essig, 1926. Ins. West. N. Amer., p. 868. —Cole, 1934. Ent.
Soc. Amer., Ann. 27: 403. — Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342: 28.
— Mallis, 1941. South. Calif. Acad. Sci., BuL 40: 91-92. —Cole, 1966. Brigham Young Univ.,
Sci. BuL, BioL Ser. 7: 19-20. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp.
112-113.




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Camponotus socius Roger
N. C. s. to Fla. w. to La.; Brazil. Ecology: Nests are built in branches and rotten
logs that are covered by sand. Introduced into the U. S.
Cam-ponotus socius Roger, 1863. Berlin. Ent. Ztschr. 7: 140. ♀.

Cam-ponotus (Tanaetnyrmex) socius var. osceola Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:
15. ♂.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 298, 319-321 (each caste).

Biology: Wheeler, 1932. N. Y. Ent. Soc, Jour. 40: 14-15. — Creighton, 1950. Harvard Univ.,
Mus. Comp. Zool., Bui. 104: 374. — Holldobler, 1971. Z. Vergl. Physiol. 75: 123-142
(recruitment behavior).

Camponotus tortuganus Emery
S. Fla. Ecology: May nest in soil under stones or under rotting wood. A
frequent house pest and reported to have been found nesting in siding and roofing of
bviildings.
Camponotus maculatus tortuganus Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 336. ♂.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 310-312 (each caste).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 13-14. —Smith, 1965. U. S. Dept. Agr.,
Tech. Bui. 1326, pp. 62-63.

Camponotus vafer Wheeler
Ariz. (Huachuca Mts., 5000 to 6000 ft.). Ecology: Nests were found under

stones.
Camponotus {Camponotus) vafer Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 315. 9,9.

Camponotus vicinus Mayr
Man., N. Dak., Colo., Okla., w. to B. C, Oreg., Calif.; Mexico. Ecology: Nests in

soil under stones or in rotting wood buried in the soil.
Formica Tejonia Buckley, 1866. Ent. Soc. Phila, Proc. 6: 161. S. Questionably the same as

vicinus.
Camponotus vicinus Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20: 940. ♂.
Camponotus (Camponotus) maculatus vicinus var. nitidiventris Emery, 1893. Zool.

Jahrb., Abt. f. System. 7: 668, 672. ♂.
Cam,ponotus (Camponotus) maculatus vicinus var. infemalis Wheeler, 1910. N. Y. Acad.

Sci., Ann. 20: 298, 305. ♂.
Camponotus {Cam.ponotus) maculatus vicinus var. luteangulus Wheeler, 1910. N. Y.

Acad. Sci., Ann. 20: 298, 304. ♀, ♂.
Cam-ponotus (CampoTVotus) maculatus vicinus var. mnritimus Wheeler, 1910. N. Y. Acad.

Sci., Ann. 20: 298, 305. $, 9, d.
Camponotus (Camponotus) maculatus vicinus var. plorabilis Wheeler, 1910. N. Y. Acad.

Sci., Ann. 20: 298, 303. 5, 9, cJ.
Camponotus (Myrmoturba) maculatus vicinus var. subrostrata Forel, 1914. Deut. Ent.

Ztschr., p. 620. ♂.
Camponotus (Myrmoturba) maculatus Maccooki berkeleyensis Forel, 1914. Deut. Ent.

Ztschr., p. 619. ♂.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 387. —Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bui. 104: 381. — Snelling, 1970. Ent. Soc. Wash., Proc. 72: 390-397. —Wheeler
and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 220 (larva).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., BuL 22: 345. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 599-560. — MaUis, 1941. South. Calif. Acad. Sci., Bui. 40: 92.
— McClure, 1943. EcoL Monog. 13: 19. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp.
170-171. —Gregg, 1963. Ants of Colo., pp. 671-675. —Cole, 1966. Brigham Young Univ., Sci.
Bui., Biol. Ser. 7: 20 (Nev. Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon,
pp. 113-114.

Genus CAMPONOTUS Subgenus MYRMOTHREX Forel

Myrmothrix Forel, 1912. Soc. Ent. de Belg., Mem. 20: 91.

Type-species: Formica abdominalis Fabricius. Desig. by Wheeler, 1913.

Most members of this subgenus are Neotropical with two forms extending into southern
United States.

Revision: Santschi, 1936. Rev. de Ent. 6: 207-218.




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abdominalis floridanus (Buckley). N. C. s. to Fla. w. to s. Miss. Ecology: Nests are in and
under rotten logs and stumps, usually in damp situations. A house infesting ant,
especially in Florida, where it is known to nest in the woodwork of buildings and feed
on household foods. May also damage beehives. C. (Af.) abdominalis abdominalis (F.)
occurs in Central and South America. Florida carpenter ant.

Formica Floridana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 161. ♂.

Camponotus atriceps stirps Yankee Forel, 1884. Soc. Vaud. des Sci. Nat., Bui. 20: 340. ♂.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 668. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 299, 325-326. — Santschi, 1936. Rev. de Ent. 6: 213-214.

Biology: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 326. —Smith, 1965. U. S. Dept. Agr., Tech.
Bui. 1326, pp. 69-70 (economic importance).
abdominalis transvectus Wheeler. S. Tex.; Mexico.

Cam,ponotus {Cam.ponotus) abdominalis transvectus Wheeler, 1910. N. Y. Acad. Sci., Ann.
20: 299,326. 5, 9, cJ.

Taxonomy: Santschi, 1936. Rev. de Ent. 6: 213.

Genus CAMPONOTUS Subgenus MYRMENTOMA Forel

MyrmentoTna Forel, 1912. Soc. Ent. de Belg., Mem. 20: 92.

Type-species: Formica lateralis Olivier. Desig. by Wheeler, 1913.

The Nearctic forms of this subgenus nest in insect galls, in branches and stems of plants,
under bark of trees, in wood and buildings, and sometimes in the soU. Colonies are small, consist-
ing of a few dozen to several hundred individuals. The listing below essentially follows that of
Creighton (1950) though there will undoubtedly be some changes in the near future.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 216-232. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool, Bui. 104: 382-390.

Camponotus anthrax Wheeler
S. Calif. Ecology: Nests were found in soil under large stones.
Camponotiis anthrax Wheeler, 1911. N. Y. Ent. Soc., Jour. 19: 96. 5, 9, 6.

Taxonomy: Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 650 (larva).

Biology: Wheeler, 1917. Amer. Acad. Arts and ScL, Proc. 52: 558. — Mallis, 1941. South. Calif.
Acad. Sci., Bui. 40: 89.

Camponotus caryae
***authority mismatch
caryae (Fitch). N. Y. s. to D. C. w. to Mich., Ohio. Ecology: Apparently associated with
trees, especially hickory.
Formica caryae Fitch, 1855. N. Y. State Agr. Soc., Trans. 14: 855. 5, 9, S.
Camponotus marginatum discolor var. cnemidatus Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 669, 678. ♂.

Taxonomy: Smith, 1940. Ent. Soc. Wash., Proc. 42: 137-141 (each caste; biol. note).

Camponotus caryae
***authority mismatch
clarithorax Emery. S. Calif.

Camponotus marginatus discolor var. clarithorax Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 670, 678. 5, 9, cJ.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 231-232 (each caste). —Wheeler,
1910. N. Y. Acad. Sci., Ann. 20: 301.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 89.

Camponotus caryae
***authority mismatch
discolor (Buckley). Ohio, S. C, Fla. w. to N. Dak., Iowa, Kans., Tex. Ecology: Nests are
in plant cavities in twigs, branches, under bark, in logs and stumps, or in insect galls. A
house infesting ant that may nest in woodwork in houses, especially in preformed
cavities or in rotting or faulty wood.
Formica discolor Buckley, 1866. Ent. Soc. Phila., Proc. 6: 166. ♂, ♀.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 230-231 (each caste). — Buren, 1944.
Iowa State Col., Jour. Sci. 18: 293.

Biology: Wheeler, 1902. Tex. Acad. Sci., Trans. 4: 23. —Wesson and Wesson, 1940. Amer.
Midland Nat. 24: 103. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 172-173. —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 70-71.




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Camponotus essigi Smith
Idaho, Oreg., Nev., Calif.

Camponotus caryae var. essigi Smith, 1923. Ent. News 24: 306. 2 .

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 89.

Camponotus hyatti
***authority mismatch
bakeri Wheeler. Calif. (Channel Islands). Ecology: Nests in soil.

Camponotus hyatti var. bakeri Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 271. ♂, ♀.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 346.

Camponotus hyatti
***authority mismatch
hyatti Emery. Nev., Calif.; Mexico (Baja Calif.). Ecology: Colonies have been found in
the stem of Yucca and in soil under a dead juniper limb.
Camponotits (Camponotus) hyatti Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 669, 680.

5.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 345-346.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 19 (Nev. Test Site).

Camponotus nearcticus Emery
Ont. s. to Fla. w. to N. Dak., Colo., Tex.; B. C. s. to Idaho, Calif. Ecology:

Forms smjill colonies in dead twigs and branches, under bark of live and dead trees, in

insect galls, pine cones, and rotting logs and stumps; also in wood products such as fenci

posts and in woodwork of houses, especially the roofing.
Formica americana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 154. 5,9. Questionably the

same as nearcticus.
Camponotus (Camponotus) marginatum var. nearcticus Emery, 1893. Zool. Jahrb., Abt. f.

System. 7: 669 675. ♂, ♀.
Camponotus (Camponotus) marginatus var. minutus Emery, 1893. Zool. Jahrb., Abt. f.

System. 7: 669, 676. ♂, ♀.
Camponotus (Camporvoius) marginatus var. decipiens Emery, 1893. Zool. Jahrb., Abt. f.

System. 7: 669,676. ♂, ♀.
Camponotus fallax fallax var. pardus Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 225. ♂.

9,6.
Camponotus fallax fallax var. tanqtuiryi Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 226.

5, 9,6.
Camponotus fallax rasilis var. pavidus Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 228.

5, 9.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300. — Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 293. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 193 Oarva).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 21: 402-403. —Wheeler, 1910. N. Y. Ent.
Soc, Jour. 18: 220-221. —Davis and Bequaert, 1922. Brooklyn Ent. Soc, Bui. 17: 24.
— Kannoswki, 1959. Insectes Sociaux 6: 134. —Gregg, 1963. Ants of Colo., pp. 675, 677.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 173-174. —Smith, 1965. U. S. Dept.
Agr., Tech. Bui. 1326, pp. 72-73 (economic importance).

Camponotus nevadensis Gregg
Nev. (Fuller Lake, 3 mi. S. Verdi, 6000 ft.).

Camponotus (Myrmentoma) nevadensis Gregg, 1973. Southwest. Nat. 18: 39-43. $ , ^, 6.

Camponotus sayi Emery
N. C. s. to Fla. w. to Nebr., Kans., Colo., Ariz., Calif. Ecology: The small colonies
are found in insect galleries in wood, in twigs and branches, insect galls, under bark, in
stalks of plants, and in rotting logs and stumps. A house infesting ant which may nest in
woodwork and feed on himian foods. In most literature as rasilis Wheeler.

Camponotus (Campotwtus) sayi Emery, 1894. ZooL Jahrb., Abt. f. System. 7: 679. ♂.

Camponotus sayi var. fricolor Pergande, 1894. Calif. Acad. Sci., Proc 4: 161. $ , 9 , 6.
Preocc. in Camponotus by Latreille, 1798.

Camponotus fallax rasilis Wheeler, 1910. N. Y. Ent. Soc, Jour. 18: 222, 227. 5, 9, cJ.

Camponotus sayi califomica Emery, 1925. In Wytsman, (Jen. Ins. 183: 118. N. name for

Camponotus bicolor Pergande


Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300. —Buren, 1944. Iowa State Col,
Jour. Sci. 18: 293-294. — SnelUng, 1968. Ent. Soc. Wash., Proc 70: 355-358.

Biology: Wheeler, 1910. N. Y. Ent. Soc, Jour. 18: 228. —Dennis, 1938. Ent. Soc Amer., Ann.
31: 302-303. —Gregg, 1963. Ants of Colo., pp. 675, 677. —Smith, 1965. U. S. Dept. Agr.,
Tech. Bui. 1326, pp. 73-74 (economic importance).




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Camponotus subbarbatus Emery
N. Y. s. to N. C. w. to Mich., Iowa, Kans. Ecology: Nests are in plant
cavities, in twigs and branches, and under bark.
Caynponotus (Camponotus) marginatus subbarbatus Emery, 1893. Zool.

Jahrb., Abt. f. System. 7: 669, 676. 3, 9, d.
Camponotus (Campotiotiis) marginatus subbarbatus var. paucipilis Emery, 1893. Zool.
Jahrb., Abt. f. System. 7: 669, 677. 5, cJ.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc, Jour. 18: 222, 229. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 300. — Buren, 1944. Iowa State Col., Jour. Sci. 18: 293.

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24: 90, 103.

Genus CAMPONOTUS Subgenus COLOBOPSIS Mayr

Colobopsis Mayr, 1861. Die Europaischen Formiciden, pp. 25, 38.

Type-species: Formica truticata Spinola. Desig. by Bingham, 1903.

Ants of this subgenus are most abundant in the southern portions of the United States. They
make their nests in hollow twigs or branches of trees and shrubs, in insect galls and nuts. The
soldiers and females have a peculiar cylindrical, truncated head which the soldiers use for
blocking the single entrance hole to the nest.

Revision: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 139-158.

Taxonomy: Wheeler and Wheeler 1953. Ent. Soc. Amer., Ann. 46: 188 (larvae).

Camponotus etiolatus Wheeler
Tex.; Mexico. Ecology: Nests have been found in insect galls and in twigs
of trees.
Camponotus (Colobopsis) abditus var. etiolatus Wheeler, 1904. Amer. Mus. Nat. Hist., Bui.
20: 150. 5, 'U, 9, 6.

Taxonomy: Wheeler, 1934. Harvard Univ., Mus. Comp. Zool., Bui. 77: 216. —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 650 (larva).

Biology: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 352-353.
hunter! Wheeler. Tex. (Victoria). Ecology: Type series taken from twig of a pecan.

Camponotus {Colobopsis) pylartes var. hunteri Wheeler, 1910. N. Y. Acad. Sci., Ann. 20:
301,353. 5, y.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 562.

Camponotus impressus (Roger)
Md. s. to Fla. w. to cent. Tex. Ecology: Colonies have been found in culms
of sedges.
Colobopsis impressa Roger, 1863. Berlin. Ent. Ztschr. 7: 160. ♀.

Taxonomy: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 144-146 (each caste). —Wheeler,
1910. N. Y. Acad. Sci., Ann. 20: 302. —Smith, 1955. Brooklyn Ent. Soc., Bui. 50: 98.

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 16.

Camponotus mississippiensis Smith
Md. s. to Fla. w. to 111., Okla., La.

Camponotus (Colobopsis) mississippietisis Smith, 1923. Psyche 30: 83. 9 , ^.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 188-190 (larva).
Biology: Smith, 1923. Ent. News 35: 127. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 303, 306.

Camponotus obliquus Smith
Ala., Miss. Ecology: Found nesting in a hickory nut.

Camponotus (Colobopsis) obliquus Smith, 1930. Ent. Soc. Amer., Ann. 23: 256. %.

Biology: Smith, 1931. Ent. News 42: 23.

Camponotus papago Creighton
Ariz.; Mexico.

Camponotus (Colobopsis) papago Creighton, 1952. Psyche 59: 153-162. 9 , "y, 9 , cJ .

Camponotus pylartes
***authority mismatch
fraxinicola Smith. N. C. to Ala. w. to Ark.

Camponotus (Colobopsis) pylartes fraxinicola Smith, 1923. Psyche 30: 86. 9, 1|.

Biology: Smith, 1924. Ent. News 35: 127.



1434 Hyjnenoptera in America North of Mexico

Camponotus pylartes
***authority mismatch
pylartes Wheeler. La., Tex. Ecology: Colonies found in twigs and spines of trees and
shrubs.
Camponotiis (Colobopsis) pylartes Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 147. ♂.

■y, 9.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 120. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 301.

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 153-158.

Genus CAMPONOTUS Subgenus MYRMAPHAENUS Emery

MyrmaphaemLS Emery, 1920. Rev. Zool. Bot. Africaines 8: 237.

Type-species: Camponotus leydigi Forel. Orig. desig.
Paracolobopsis Emery, 1920. Rev. Zool. Bot. Africaines 8: 249.

Type-sf>ecies: Camponotus salvini Forel. Orig. desig.
Neomyrmamblys Wheeler, 1921. Psyche 28: 19.

Type-species: Camponotus fastigatus Roger. Desig. by Santschi, 1921.

Only two species of this Neotropical subgenus reach the United States.

Camponotus ulcerosus Wheeler
Tex., Ariz.; Mexico. Ecology: Nests are constructed in the soil beneath
stones. A carton shield is constructed at the nest entrance with an opening the same
size as the head of the major worker. Thus, the major worker functions as a door in this
opening.

Camponotus {Cam,ponotus) ulcerosus Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 351. ♂.

Camponotus (Campmiotus) bruesi Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 349. ♂.

Taxonomy: Creighton, 1951. Psyche 58: 47-64 (also biology).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 562. —Creighton, 1953. Psyche
60: 82-84. — Samuelson, 1961. Pan-Pacific Ent. 37: 189 (association with larvae of Strymon
melinus).

Camponotus yogi Wheeler
S. Calif. Ecology: Most colonies have been found in living stems of Haplopappfus
pinifolius where they live in burrows made by buprestids.
Camponotus yogi Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 420. ♂.

Taxonomy: Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 650 (larva). — Creighton
and Snelling, 1966. Psyche 73: 187-195 (female, male; biological notes).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 562. — Mallis, 1941. South. Calif.
Acad. Sci., Bui. 40: 92.

Genus CAMPONOTUS Subgenus MYRMOBRACHYS Forel

Myrmobrachys Forel, 1912. Soc. Ent. de Belg., Mem. 20: 91.

Type-species: Formica senex Smith. Desig. by Wheeler, 1913.

This Neotropical subgenus extends into Florida, Texas, and Arizona. The species found in the
United States usually nest under bark of trees, in branches of trees and shrubs, and in logs and
stumps.

Camponotus mina Forel
S. Ariz.; Mexico. Ecology: Most colonies have been found nesting in mesquite.
Camponotus Mina Forel, 1879. Soc. Vaud. des Sci. Nat, Bui. 16: 83. ♂.
Camponotus erythroprus Pergande, 1894. Calif. Acad. Sci., Proc. 4: 28. ♀.
Camponotus (Camponotus) mina zuni Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300. ♂.

Taxonomy: Creighton, 1965. Amer. Mus. Novitates 2239: 1-7 (also biological notes).

Camponotus planatus Roger
S. Fla., S. Tex.; W. Indies, Mexico. Ecology: Arboreal, nesting in branches and
under bark.
Camponotus planatus Roger, 1863. Berlin. Ent. Ztschr. 7: 148. 5, 9, 6.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 301, 348-349. —Wheeler and Wheeler,
1953. Ent. Soc. Amer., Ann. 46: 194 (larva).



Superfamily FORMICOIDEA 1435

Biology: Wheeler, 1932. N. Y. Ent. Soc, Jour. 40: 15. —Wheeler, 1942. Harvard Univ., Mus.
Comp. Zool., Bui. 90: 258.

Camponotus trepidulus Creighton
S. Ariz. (Baboquivari Mtns.). Ecology: Colonies were found in dead limbs
of Quercus oblongifolia.
Camponotus {Myrmobrachys) trepidulus Creighton, 1965. Amer. Mus. Novitates 2239: 7-9.
^,'U, 9, 6.

Tribe LASIINI

Genus LASIUS Fabricius

Ants of this holarctic genus nest in exposed soil, under objects or in rotting wood. Colonies are
small to moderate in size. Workers attend and may foster honeydew-excreting insects. The
treatment here follows the revision by Wilson (1955).

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637-639. —Wheeler, 1916. Psyche 23:
168-173. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 1-199.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 147-150 (larvae).

Biology: Kannowski, 1959. Insectes Sociaux 6: 145-146 (flight comparisons). — Pontin, 1961.
Jour. Animal Ecology 30: 47-54 (population stabilization and competition between L. flavus
and L. niger). —Pontin, 1963. Jour. Animal Ecology 32: 565-567 (competition and ecology
of L. flavus and L. niger).

Morphology: Bemardi, et. al., 1967. Tetrahedron Letters 40: 3893-3896 (components of
secretion of mandibular glands).

Genus LASIUS Subgenus LASIUS Fabricius

Lasius Fabricius, 1805. Systema Piezatorum, p. 415.

Type-species: Formica nigra Linnaeus. Desig. by Bingham, 1903.

Donistharpea Morice and Durrant, 1914. Ent. Soc. London, Trans., pp. 421-423.
Type-species: Formica nigra Linnaeus. Orig. desig.

Lasius alienus (Foerster)
N. S., N. B. s. to Fla. w. to Man. N. Dak, S. Dak., Neb., Kans., Ark., Miss.;
B. C, Mont., Idaho, Wash., Oreg., Calif., S. Ariz.; Mexico; Eurasia. Ecology: In N. Amer.,
this ant shows a preference for well-shaded woodlands where it nests in rotting logs
and stumps or under stones. Only occasionally has it been found in open areas. In the
early American literature, this species has been recorded as americanus Emery, and

Lasius alienus
***authority mismatch
has commonly been confused with neoniger. A frequent house pest. Cornfield
ant is the approved conunon name for alienus.

Formica aliena Foerster, 1850. Hym. Studien 1: 36-38. 5, cJ.

Prenolepis Uisioides Emery, 1869. Accad. Natur. Napoli, Ann. 2: 6-7. 5, 9, 6.

Prenolepis fuscula Emery, 1869. Accad. Natur. Napoli, Ann. 2: 8. ♂.

Lasius fumatus Emery, 1870. Soc. Ent. Ital., Bol. 2: 194.

Lasius niger var. americanus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 639. 9 , 9, 6.

Lasius niger var. grandis Forel, 1909. Soc. Ent. de Belg., Ann. 53: 104-105. ♂.

Lasius niger alienus var. alieno-americanus Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 525-526. ♀.

Lasius niger turcicus Santschi, 1921. Soc. Esp. Hist. Nat., Bol. 21: 115-116. ♂ (♀ misdet.).

Lasius niger lasioides var. barbara Santschi, 1921. Soc. Esp. Hist. Nat., Bol. 21: 170. ♀.

Acanthomyops niger alienus var.flavidus Kuznetzov-Ugamskij, 1927. Rev. Russ. d'Ent.
21: 189. ♂.

Acanthomyops niger alienus var. turkmenus Kuznetzov-Ugamskij, 1927. Rev. Russ. d'Ent.

21: 189. ♂.
Lasius brunneus var. obscurata Stitz, 1930. Mitt. Zool. Mus. Berlin 16: 239-240. ♂, ♀.
Lasius alienus illyncus Zimmermann, 1934. Zool.-Bot. GcseU. Wien, Verb. 84: 50-52. ♂, ♀.

6.
Lasius alienus var. pannonica Roszler, 1942. Siebenburgischer Verb. Naturw.,

Hermannstadt, Verb, und Mitt. 91-92 :40. ♂, ♀.
Lasius alienus var. pontica Starcke, 1944. Ent. Ber. 11: 156-157. ♂.




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Taxonomy: Gregg, 1945. Ent. Soc. Amer., Ann. 38: 529-546. —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 46: 147-148 (larva).

Biology: Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 77-89. —Benjamin, 1958.
Canad. Ent. 90: 419. — Kannowski, 1959. Insectes Sociaux 6: 135-136. —Wheeler and
Wheeler, 1963. Ants of N. Dak., pp. 178-179. —Gregg, 1963. Ants of Colo., pp. 454-457.
—Bums, 1964. Ent. Soc. Amer., Ann. 57: 138 (association with tuliptree scale). — Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 80-81 (economic importance). —Wheeler and
Wheeler, 1973. Ants of Deep Canyon, pp. 115-116.

Morphology: Regnier and Wilson, 1969. Jour. Ins. Physiology 15: 893-898 (alarm-defense
system).

Lasius crypticus Wilson
N. Dak. s. to N. Mex. w. to Alta., Idaho, Oreg., Utah; S. Calif. Ecology:

Abundant in prairie regions where it nests under stones or occasionally in crater nests
in soil.
Lasius {Lasius) crypticus Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113:
104-118. 5, 9, d.

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 179-182.

Lasius neoniger Emery
Que., Maine s. to Fla. w. to Idaho, Wyo., Colo., N. Mex.; Calif. (Sierras);

Alaska (?). Ecology: Nests almost exclusively in open areas, either under stones or in
crater nests. One of the dominant ants found in lawns, cultivated fields, grassy road
strips, and prairies. A common house and lawn pest and also fosters honeydew-excreting
insects. In American literature, neoniger has often been confused with alienus
i=americamis Emery), and many early references applying these names are here
referred to neoniger.
Lasius niger var. neoniger Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637. ♂.

Taxonomy: Gregg, 1945. Ent. Soc. Amer., Ann. 38: 534.

Biology: Forbes, 1894. 18th Rpt. State Ent. 111., 171 pp. —Forbes, 1908. 111. Agr. Expt. Sta.
Bui. 131: 31-44. — Tanquary, 1913. lU. State Lab., Nat. Hist. Bui. 9: 417-143. — Metcalf and
Flint, 1939. Destructive and Useful Insects, pp. 371-374, 770. — Severin, 1920. S. Dak. State
Ent. Cir. 20: 3. —Talbot, 1945. Amer. Midland Nat. 34: 504-506. —Talbot, 1946. Ecology 27:
65-70. — Schread and Chapman, 1948. Conn. (State) Agr. Expt. Sta. Bui. 515: 4-11.
—Talbot, 1953. Mich. Univ., Lab. Vertebrate Biol., Contrib. No. 63, pp. 3-12. —Wilson,
1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 100-104. — Ayre, 1962. Canad. Jour. Zool.
40: 157-164 (parasitism). —Wheeler and Wheeler, 1963. Ants, of N. Dak., pp. 182-185.
—Gregg, 1963. Ants of Colo., pp. 461^65. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326,
pp. 81-84 (economic importance).

Lasius niger (Linnaeus)
Colo., N. Mex., Ariz., Utah, Mont., Idaho, Wash., Oreg., Calif.; Mexico;

Eurasia. Ecology: Found in forests or in open situations where it usually nests under

stones, though occasionally in rotting wood.
Formica nigra Linnaeus, 1758. Syst. Nat., Ed. 10, 1: 580. 2 .

Lasius niger var. alieno-niger Forel, 1874. Les Fourmis de la Suisse, pp. 47, 49. ♂, ♀.
Lasius niger var. alienoides Emery, 1891. Explor. Sci. Tunisie, Paris, p. 16. ♂.
Lasius niger fiavescens Forel, 1903. Mus. Zool. Acad. Imp. Sci. St. Petersburg, Ann. 8:

386-387. ♂.
Lasius niger emeryi Ruzsky, 1905. Schrift. Naturforsch.-Ges. Univ. Kasan 38: 313-314. ♂.
Acanthomyops niger nitidus Kuznetzov-Ugamskij, 1927. Rev. Russ. d'Ent. 21: 188. ♂.
Acanthomyops niger alienus var. pilicomis Kuznetzov-Ugamskij, 1927. Rev. Russe d'Ent.

21: 189. ♀.
Acantfiomyops niger var. minimus Kuznetzov-Ugamskij, 1928. Ants of South Ussuri

Region, U. S. S. R. Natl. Geog. Soc. Publ., p. 20. ♂.
Lasius emarginatu^ var. nigrescens Stitz, 1930. Mitt. Zool. Mus. Berlin 16: 240. ♀.
Lasius niger colcrratus Santschi, 1937 Soc. Ent. de Belg., Bui. 68: 387. ♂, ♀.
Lasius emarginatus vbt. japonicus Santschi, 1941. Mitt. Schweiz. Ent. Ges., 18: 277-278. ♂.

9.
Lasius transylvanica Roszler, 1943. Zool. Anz. 144: 44-46. 5, cJ.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 148-150 (larva).




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Biology: Goswald, 1932. Ztschr. Wiss. Zool. 142: 1-156 (in Germany). —Wilson, 1955. Harvard
Univ., Mus. Comp. Zool., Bui. 113: 72-75. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, pp. 116-117.

Lasius pallitarsis (Provancher)
N. S., Que. w. to B. C, Alaska s. to N. Y., N. C, Mich., Wis., Minn., S.
Dak., N. Mex., Ariz., Nev., Calif.; Siberia. Ecology: Found mostly in forested areas
where it nests in rotting logs and stumps or under stones. Occasionally a house infesting
ant.
Formica pallitarsis Provancher, 1881. Nat. Canad. 12: 355. 5, d.
Lasius niger sitkaensis Pergande, 1900. Wash. Acad. Sci., Proc. 2: 519. ♂.

Taxonomy: Wheeler, 1917. Harvard Univ., Mus. Comp. Zool, Bui. 61: 18, 21. —Cole, 1942.
Amer. Midland Nat. 28: 374. — Francoeur and Beique, 1966. Canad. Ent. 98: 144
(Provancher types).

Biology: Wheeler, 1915. Psyche 22: 206. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool.,
Bui. 113: 45-47. — Medler, 1958. Ent. Soc. Wash., Proc. 60: 258 (flights). —Wheeler and
Wheeler, 1963. Ants of N. Dak., pp. 185-188. —Gregg, 1963. Ants of Colo., pp. 461-465.
—Corbet and Ayre, 1968. Canad. Field-Nat. 82: 230-231 (swarming and mating).
— Kannowski, 1969. Canad. Field-Nat. 83: 283-285 (nuptial flights). —Corbet and Ayre,
1969. Canad. Field-Nat. 83: 285-286 (swarming and mating). — Akre and HiU, 1973. Kans.
Ent. Soc, Jour. 46: 526-536 (behavior of a myrmecophilous beetle).

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome numbers).

Lasius sitiens Wilson
Colo., N. Mex., Ariz., Nev.; Mexico. Ecology: Found in the lower altitudinal
forest belts in mountains, 7000 to 8000 feet where it nests under stones in dry open
situations.
Lasius (Lasius) sitiens Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 108-111.
5, 9,6.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 20 (Nev. Test Site).

Genus LASIUS Subgenus CAUTOLASIUS Wilson

Lasiiis subg. Cautolasius Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 13.
Type-species: Formica flava Fabricius. Orig. desig.

Lasius fallax Wilson
Mont., Wyo., Colo., Idaho, Utah, Ariz., Wash. Ecology: Colonies have been found

under stones in forest clearings.
Lasius (Cautolasius) fallax Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 1 13:

130-133. 5, 9,6.

Lasius flavus (Fabricius)
N. S., N. B., Que. s. to Ala. w. to Alta., Wash., Oreg., Calif.; Eurasia.

Ecology: A subterranean ant which nests in various situations but most often under

stones. Known to build mounds in parts of Eurasia. Workers may attend aphids on roots

of grasses.
Formica flava Fabricius, 1781. Species Insectorum 1: 491. ♂.
Lasius brevicomis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637. 9, 9, S.
Lasius flavusmyops Forel, 1894. Soc. Vaud. des Sci. Nat., Bui. 30: 12. ♀.
Lasius flavus myops var. Jlavoides Forel, 1894. Soc. Vaud. des Sci. Nat., Bui. 30: 12. ♂.
Lasius flavus var. fuscoides Ruzsky, 1902. Schrift. Naturforsch. CJesell. Univ. Kasan 38:

281.
La^us flavus var. odoratus Ruzsky, 1902. Schrift. Naturforsch. Gesell. Univ. Kasan 38:

282-283. ♂.
Lasius flavus var. flavo-myops Forel, 1915. Mitt. Schweiz. Ent. Gesell. 12: 52.
Lasius iFormicina) brevicomis microps Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.

52: 526. ♂.
Lasius iFormicina) flavus claripennis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:

527. 5, 9, S.
Formicina flava var. morbosa Bondroit, 1918. Soc. Ent. de France, Ann. 87: 28-29. 9,9.
Lasius umbratus var. apennina Menozzi, 1924. Atti Soc. Nat. Mat. Modena 8: 15. 2 .
Lasius umbratus ibericus Santschi, 1925. Eos 1: 349-350. ♂.
Lasius umbratus ibericus var. sancho Santschi, 1925. Eos 1: 350. ♂.
Lasius flavus var. olivacea Karawajew, 1926. Konowia 5: 194. ♀.




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Lasius iChthonolasius) helveolus Cook, 1950. Ants of Calif., p. 327. ♂.
Lasius (Chthonolasius) helmis Cook, 1950. Ants of Calif., p. 326. 5? . In figs.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24: 68-69. — Buren, 1944. Iowa State Col., Jour.
Sci. 18: 296. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 152-154 (larva).
—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 396. — Goswald, 1932. Ztschr. Wiss.
Zool. 142: 1-156 (in Germany). —Wesson and Wesson, 1940. Amer. Midland Nat. 24: 100.
—Cole, 1942. Amer. Midland Nat. 28: 375. —Wheeler and Wheeler, 1944. N. Dak. Hist.
Quart. 11: 253. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 126-128.
— Waloff, 1957. Insectes Sociaux 4: 391-408 (effect of number of queens on their survival
and the development of first brood). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
189-191. —Gregg, 1963. Ants of Colo., pp. 465-469. — Marikovsky, 1965. Insectes Sociaux
12: 63. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 117.

Lasius nearcticus Wheeler
Que., Ont. s. to N. C, Tenn. w. to S. Dak., Wyo., Colo. Ecology:

Subterranean; prefers dense, moist woodlands here it nests in soil under rocks or fallen

logs.
Formica mellea Provancher, 1881. Nat. Canad. 12: 356. ♂. Preocc. by Say, 1836.
LasiiLS flavtts nearcticus Wheeler, 1906. Psyche 13: 38. ♂.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24: 68, 70. —Buren, 1944. Iowa State Col., Jour.
Sci. 18: 296, 297. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 151-152 (larva).
— Francoeur and Beique, 1966. Canad. Ent. 98: 144 (Provancher types).

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24: 100. — Headley, 1943. Ohio Jour.
Sci. 43: 29. —Hicks, 1947. Canad. Ent. 79: 170-171 (imusual nesting site). —Wilson, 1955.
Harvard Univ., Mus. Comp. Zool., Bui. 113: 135-136. —Gregg, 1963. Ants of Colo., pp.
469-471.

Genus LASIUS Subgenus CHTHONOLASIUS Ruzsky

Chthonolasius Ruzsky, 1913. Arch. f. Naturgesch. 79: 59-61.

Type-species: Formica umbrata Nylander. Desig. by Emery, 1925.

Ants of this subgenus are more subterranean than those of the subgenus Lasius, and some
build large earthen moimds. Food is largely honeydew derived from subterranean plant lice and
mealybugs. Some forms are temporary parasites on ants of the subgenus Lasius.

Lasius atopus Cole
Calif. (3 mi. S. Leggett, Mendocino Co.). Ecology: The nest was found in dry soil
under a stone in an unshaded area.
Lasius (Chthonolasius) atopus Cole, 1958. Tenn. Acad. Sci., Jour. 33: 75-77. ♂.

Lasius humilis Wheeler
Colo., N. Mex., Nev. Ecology: Colonies were found under stones in moist soil
of a meadow and in moist open woods.
Lasius (Farmicina) humilis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 528. ♂, ♀.

Biology: Gregg, 1963. Ants of Colo., pp. 473^75.

Lasius minutus Emery
N. S., Maine s. to Va. w. to Minn., Iowa. Ecology: Most often foui.d in

sphagnum bogs, swampy meadows, or open dry forests. Taken most often in mounds or
masonry domes. Some specimens have been associated with L. alienus.
Lasius umbratus minutus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 37, 641. 5, 9, 6.

Taxonomy: Wheeler, 1910. Psyche 17: 237, 238, 241-242.

Biology: Gaige, 1914. Univ. Mich., Mus. Zool., Occas. Papers 5: 3, 4, 21, 23. —Wheeler, 1915.
Psyche 22: 206. -Morris, 1943. Ind. Acad. Sci., Proc. 52: 215. — WUson, 1955. Harvard
Univ., Mus. Comp. Zool., BuL 113: 182. — Kannowski, 1959. Insectes Sociaux 6: 136-138,
151-153 (colony founding; possible parasitism on Lasius alienus and L. pallitarsis (=
sitkaensis)). —Kannowski, 1959. Ecology 40: 162-165 (radioactive phosphorus in study of
colony distribution)




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Lasius nevadensis Cole
Nev. (Kyle Canyon, Charleston Mts.). Ecology: Nests were found in an

unshaded area in open forests; some were under stones and some were in exposed soil
with a scattering of soil around the entrance.
Lasius iChthonolasius) nevadensis Cole, 1956. Tenn. Acad. ScL, Jour. 31: 26-27. 5, 9, 6.

Lasius speculiventris Emery
N. J., Pa. w. to Minn., Iowa, Kans. Ecology: Nests have been found
under rocks and in rotting wood either in pastures or wooded areas.
Lasius speculiventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637, 641. 5, cJ.

Taxonomy: Wheeler, 1910. Psyche 17: 237, 242-243. —Wheeler, 1917. Conn. State Geol. and
Nat. Hist. Survey Bui. 22: 592.

Biology: Wilson, 1955. Harvard Univ., Mus. Comp. Zool, Bui. 113: 173. — Kannowski, 1959.
Insectes Sociaux 6: 138-141, 153-154 (parasitism on minutus and possible other Lasius
spp.).

Lasius subumbratus Viereck
N. S., Maine w. to Sask., Wash., Oreg., s. to N. Mex., Ariz., Nev.

Ecology: Nests under stones or rotting logs in meadows and forests. A temporary social
parasite. Host: Lasius pallitarsis (Provancher).
Lasius umbratus subumbratus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 73. ♀.

Taxonomy: Wheeler, 1910. Psyche 17: 237-239. —Cole, 1942. Amer. Midland Nat. 28: 374.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 528. —Wheeler, 1917. Psyche
24: 167-176. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 179-180. —Wheeler
and Wheeler, 1963. Ants of N. Dak, pp. 191-192. —Gregg, 1963. Ants of Colo., pp. 475-478.

Lasius umbratus (Nylander)
N. S., N. B., Que. s. to Fla. w. to Idaho, Utah, Ariz.; Eiu-asia. Ecology:

Prefers moist soil and most commonly nests under stones and in or imder rotting logs

and stumps. May foster subterranean plant lice and mealybugs; occasionally a

house-infesting ant. A temporary socicd parasite. Host: Lasius alienus (Foerster), L.

Lasius niger
***authority mismatch
(L.), L. neoniger Emery (?).
Formica umbrata Nylander, 1846. Acta. Soc. Sci. Fenn. 2: 1048-1050. ♀, ♂.
Formica mixta Nylander, 1846. Acta. Soc. Sci. Fenn. 2: 1050-1052. ♀.
Formica affinis Schenck, 1852. Jahrb. Ver. Nat. Nassau 8: 62-63. 5, 9, S.
Formica aphidicola Walsh, 1862. Ent. Soc. Phila., Proc. 1: 310. 5, 6.
Lasuis umbratus var. mixto-umbratus Forel, 1874. Nouv. Mem. Soc. Helv. Sci. Nat., p. 48.

5.
Lasius umbratus var. exacutus Ruzsky, 1904. Kasan Univ. Obschchestvo

estestvoispytatelei Protokoly Zasiedanii, no. 206, p. 15. ♂.
Lasius umbratus var. mixto-affinis Ruzsky, 1904. Kasan Univ. Obschchestvo

estestvoispytatelei Protokoly Zasiedanii, no. 206, p. 15. Nomen nudum.
Lasius umbratus var. mixto-bicomis Ruzsky, 1905. Schrift. Naturforsch. (Jesell. Univ.

Kasan 38: 292. Nomen nudum.
Lasius umbratus var. affino-um,bratvs Donisthorpe, 1914. Ent. Rec. 26: 40. ♂.
Lasius umbratus var. jrrzewalskii Ruzsky, 1915. Mus. Zool. Acad. Sci. Petrograd 20: 434.

5.
Formicina umbrata distinguenda Emery, 1916. Rend. Accad. Bologna, pp. 64-65. ♂, ♀.
Formicina umbrata var. hybrida Emery, 1916. Rend. Accad. Bologna, p. 66.
Formicina umbrata var. nuda Bondroit, 1917. Soc. Ent. de France, Bui. 86: 176.
Formicina umbrata var. sabularum Bondroit, 1918. Soc. Ent. de France, Bui. 87: 31.
Formicina belgarum Bondroit, 1918. Soc. Ent. de France, BuL 87: 31. ♂, ♀.
Lasius bicomis var. citrina Emery, 1922. Soc. Ent. Ital, Bol. 54: 12. ♂.
Lasius umbratus var. viehmeyeri Emery, 1922. Soc. Ent. Ital., Bol. 54: 13-15. ♂, ♀.
Lasius silvestrii Wheeler, 1928. Lab. Zool. Portici, Bol. 20: 120-121. ♀.
Lasius viehmeyeri var. dalmatica Starcke, 1937. Tijdschr. Ent. 80: 53-54. ♀.
Lasius umbratus var. hirtiscapus Starcke, 1937. Tijdschr. Ent. 80: 43. ♀.
Lasius umbratus distinguendus var. cereomicans Starcke, 1937. Tijdschr. Ent. 80: 48-49.

9, 9, 6.
Lasius silvestri (!) var. osakana Santschi, 1941. Mitt. Schweiz. Ent. Ges. 18: 278. ♀.
ChtonolasiusC) affinis var. nyaradi Roszler, 1943. Zool. Anz. 144: 47-48. ♂, ♀.
Lasius umbratus epinotalis Buren, 1944. Iowa State Col., Jour. ScL 18: 296, 297. ♂.




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Taxonomy: Wheeler, 1910. Psyche 17: 237-241. —Wheeler, 1917. Conn. State Geol. and Nat.
Hist. Survey Bui. 22: 592. —Wheeler "and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 150-151
(larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 397. —Smith, 1928. Ent. News 39:
277-278. —Wesson and Wesson, 1940. Amer. Midland Nat. 24: 90, 101. —Wilson, 1955.
Harvard Univ., Mus. Comp. ZooL, Bui. 113: 161-165. — Kannowski, 1959. Insectes Sociaux
6: 141, 154-155 (possible parasitism on other Lasitis spp.). — Wheeler and Wheeler, 1963.
Ants of N. Dak, pp. 192-194. —Gregg, 1963. Ants of Colo., pp. 478-481. —Smith, 1965. U.
S. Dept. Agr., Tech. Bui. 1326, pp. 84-86 (economic importance).

Morphology: Blum, et. al, 1968. Ent. Soc. Amer., Ann. 61: 1354-1359 (terpenes in mandibular
glands). — Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).

Lasius vestitus Wheeler
B. C, Idaho, Oreg., Calif.

Lasiiis umbratus vestitus Wheeler, 1910. Psyche 17: 238, 242. ♀.

Lasius aChthonolasius) pilostLS Smith, 1934. Ent. Soc. Amer., Ann. 27: 384. ♂.

Biology: Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bui. 113: 173-175.

Genus ACANTHOMYOPS Mayr

Acanthomyops Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 699.
Type-species: Formica clavigera Roger. Monotypic.

The ants of this exclusively North American genus nest in the soil, usually beneath objects,
and also in rotting logs and stumps. They are mostly subterranean in habit. The females and
workers have a characteristic citronella or lemon-verbena odor. Species of Acanthomyops are
sometimes of economic importance because they foster honeydew excreting insects and because
of their objectionable habit of nesting around foundations of buildings and beneath basement
floors. The vringed forms are often mistaken for termites.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 638, 642. —Wheeler, 1916. Psyche 23:
170-172. — Buren, 1950. Ent. Soc. Wash., Proc. 52: 184-190. —Wing, 1968. Cornell Univ.,
Agr. Expt. Sta., Mem. 405, 173 pp. (also biological notes for each species).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 154-156 Garvae).
—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 648 (larvae).

Biology: Kannowski, 1963. Pavia Univ., Symp. CJenet. Biol. Ital. 12: 74-102 (flight activities).
—Talbot, 1963. Ecology 44: 549-555 (local distribution and flight activities).

Acanthomyops arizonicus (Wheeler)
S. Ariz. Ecology: Most collections have been from under stones.

LasiTis (Acanthom.yops) interjectus arizoniciis Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:532. ♂.

Acanthomyops bureni Wing
Wis. (Comstock, Barron Co.).

Acanthomyops bureni Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 135-138.

califomicus (Wheeler). S. Calif. Ecology: Colonies have been found in soil under stones.

Lasius {Acanthomyops) interjectus califomicus Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:531. ♂, ♀.

Acanthomyops claviger (Roger)
Mass., Ont., N. Y. s. to Fla. w. to Minn., Nebr., Kans., Miss. Ecology: Found
in woodlands, pastures, or open fields where they nest imder stones, in rotting wood,
and occasionally in exposed soil. A common house pest. SmaUer yellow ant.
Formica clavigera Roger, 1862. Berlin. Ent. Ztschr. 6: 241. ♀.
Lasius (Acanthomyops) parvula Smith, 1934. Psyche 41: 213. §.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 296. —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 46: 155-156 Oarva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 398. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31: 296, 397. — Rau, 1945. Ent. News 56: 119. —Carter, 1962. Elisha Mitchell Sci. Soc.,
Jour. 78: 150-204. —Talbot, 1963. Ecology 44: 549-557 (local distribution, flight activities).
—Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 86-87 (economic importance). —Talbot,
1973. Great Lakes Ent. 6: 20-21 (S. Mich.).




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Morphology: Wheeler and McClendon, 1903. Biol. Bui. 4: 149-155. — Chadha, et al., 1962.
Jour. Ins. Physiol. 8: 175-179 (citronellal and citral in mandibular gland secretion).
— Regnier and Wilson, 1968. Jour. Ins. Physiol. 14: 955-970 (alarm-defense system).

Acanthomyops colei Wing
N. Mex., Ariz.

Acanthomyops colei Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 88-89. 5 , c? .

Acanthomyops coloradensis (Wheeler)
Man., N. Dak., S. Dak., Colo., N. Mex. w. to Alta., Oreg., Utah.

Ecology: Most colonies have been found under stones. Confused with A. claviger in
earlier literature.
Lasius {Acanthomyops) interjectus coloradensis Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 532. 5, 9, d.

Biology: Gregg, 1963. Ants of Colo., pp. 481, 483-484. —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 195-197 (reported as claviger).

Acanthomyops creightoni Wing
Utah (Moab, Grand Co.).

Acanthomyops creightoni Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp.
141-143. 5, 9, 6.

Acanthomyops interjectus (Mayr)
Mass., N. Y. s. to Ga. w. to Mont., Idaho, Utah, N. Mex. Ecology: Found in
woodlands, pastures or meadows. They may nest in exposed soil where the nest is
sometimes surmounted by a mound, under stones or other objects, in rotting logs and
stumps, or next to foundation walls of buildings. A frequent house pest. Larger yellow
ant.
Lasius (Acanthomyops) interjectus Mayr, 1866. Zool.-Bot. Gesell. Wien, Verh. 16: 888. ♀.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22: 592, 594.

— Buren, 1944. Iowa State Col, Jour. Sci. 18: 296. —Wheeler and Wheeler, 1953. Ent. Soc.
Amer., Ann. 46: 156 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 397-398. —Smith, 1928. Kans. Ent.
Soc, Jour. 1: 14-18. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 296, 306. —Carter, 1962.
Elisha MitcheU Sci. Soc., Jour. 88: 150-204. —Talbot, 1963. Ecology 44: 549-557 Oocal
distribution, flight activities). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 197-200.
—Gregg, 1963. Ants of Colo., pp. 484-486. —Smith, 1965. U. S. Dept. Agr., Tech. Bui. 1326,
pp. 88-89 (economic importance). —Talbot, 1973. Great Lakes Ent. 6: 22 (S. Mich.).

Acanthomyops latipes (Walsh)
Que., Maine w. to B. C. s. to S. C, Tenn., 111., Iowa, Okla, N. Mex., Ariz., Calif.
Ecology: Found in open woodlands, meadows, or pastures where they may nest in
exposed soU commonly surmounted by a mound, under stones or other objects, or at the
base of stumps.
Formica latipes Walsh, 1862. Ent. Soc. Phila., Proc. 1: 311. 5, 9, 6.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 638, 642. —Wheeler, 1917. Conn.
State Geol. and Nat. Hist. Survey Bui. 22: 592. —Buren, 1944. Iowa State Col., Jour. Sci.
18: 296. —Cole, 1954. Tenn. Acad. Sci., Jour. 29: 284. —Wing, 1968. ComeU Univ., Agr.
Expt. Sta., Mem. 405, pp. 98-105 (hybrids described).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 398. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 21: 398. — Mallis, 1941. South. Calif. Acad. Sci., Bui. 41: 79. —Talbot,
1963. Ecology 44: 549-557 (local distribution, flight activities). —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 200-202. —Gregg, 1963. Ants of Colo., pp. 48&488. —Smith,
1965. U. S. Dept. Agr., Tech. Bui. 1326, pp. 91-93 (economic importance). —Cole, 1966.
Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 20 (Nev. Test Site). —Talbot, 1973. Great
Lakes Ent. 6: 22 (S. Mich).

Morphology: Wheeler and McClendon, 1903. Biol. Bui. 4: 149-155. —Wheeler, 1903. Amer.
Mus. Nat. Hist, Bui. 41: 79.

Acanthomyops murphyi (Forel)
N. Y., Ont. s. to Ga. w. to Sask., Idaho, n. Calif., Utah, N. Mex. Ecology:

Usually nests under or next to stones in open woodlands or edges of woodlands. Prefers
sandy soil. Widely but sporadically distributed in N. Amer.
Lasius (Acanthomyops) Murphii Forel, 1901. Soc. Ent. de Belg., Ann. 45: 367. $, 9, d.
Lasius (Acanthomyops) murphyi Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 398.
Emend.




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Taxonomy: Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 13-121 (hybrids
described).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 398. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 530. —Talbot, 1963. Ecology 44: 549-557 (local distribution, flight
activities). —Gregg, 1963. Ants of Colo., pp. 489^90. —Smith, 1965. U. S. Dept. Agr., Tech.
Bui. 1326, pp. 89-91 (economic importance). —Talbot, 1973. Great Lakes Ent. 6: 21 (S.
Mich.).

Acanthomyops occidentalis (Wheeler)
Man., Minn., Nebr., Colo., N. Mex. w. to B. C, Wash., Wyo., Utah.
Ecology: Nests have been found under stones in dry sandy soil.
Lasius (.Acanthomyops) occidentalis Wheeler, 1909. N. Y. Ent. Soc, Jour. 17: 83. §, 9, 6.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 530. —Gregg, 1963. Ants of
Colo., pp. 490-493.

Acanthomyops plumopilosus (Buren)
N. C, Mich., Minn., Iowa. Ecology: Found under stones and in rotting
logs.
Lasius (Acanthomyops) plumopilosus Buren, 1941. Iowa State Col., Jour. Sci. 15: 231-235.
5, 9, d.

Biology: Buren, 1944. Iowa State Col., Jour. Sci. 18: 296, 299.
pogonog:ynus (Buren). Iowa, Colo., Idaho. Listed as a hybrid of m,urphyi x latipes by Wing
(1968).
Lasius {Acanthomyops) pogonogynu^ Buren, 1950. Ent. Soc. Wash., Proc. 52: 186. ♂, ♀.

Taxonomy: Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 117-119 (as a hybrid).
Biology: Gregg, 1963. Ants of Colo., pp. 493-494.

Acanthomyops pubescens (Buren)
Minn. Ecology: Nests with low mounds were found in soil in open woods.
Lasius {Acanthomyops) pubescens Buren, 1942. Iowa State Col., Jour. Sci. 16: 405. ♂, ♀.

Acanthomyops subglaber (Emery)
Maine s. to Ga., Tenn. w. to Sask., N. Dak., S. Dak. Ecology: Nests in

woodlands or open areas, in mound nests, under stones, or in or under rotting logs and
stumps.
Lasius claviger var. subglaber Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 642. 5, 9, 6.
Lasius {Acanthomyops) clavigeroides Buren, 1942. Iowa State Col., Jour. Sci. 16: 406. ♀.
9, cJ.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist, Bui. 21: 398. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 533. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 155
(larva). —Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 121-132 (hybrids
described). —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 648, 649 (larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 623. —Wheeler and Wheeler, 1963.
Ants of N. Dak, pp. 203-204. —Talbot, 1973. Great Lakes Ent. 6: 19-20 (S. Mich.).

Genus PARATRECHINA Motschulsky

Species of this genus often nest in the soil where the entrance may be surrounded by a small
irregular crater of earth, under stones, or beneath moss. They are occasionally house pests.
Because of the differences of opinion in recent literature regarding the status of subgenera, par-
ticularly Nylanderia, the two subgenera are listed as in previous catalogs.

Genus PARATRECHINA Subgenus PARATRECHINA Motschulsky

Paratrechina Motschulsky, 1863. Soc. Nat. Moscou, Bui. 36: 13.

Type-species: Paratrechina currens Motschulsky. Desig. by Wheeler, 1911.

longicomis (Latreille). S. C. s. to Fla. w. to Tex.; Calif.; Pantropical. Ecology: Highly adaptable
and may nest in many situations such as trash, refuse, plant cavities, rotting wood, and
in soil under stones. Workers are omnivorous and feed on live and dead insects, seeds,
honeydew and household foods. Sometimes a household pest. Introduced into the U. S.
Probably native to Africa or the Orient and spread to many parts of the world by




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commerce. Sometimes found in northern states in greenhouses or other buildings. Crazy

ant.
Formica longicomis Latreille, 1802. Hist. Nat Fourmis, p. 113. ♂.
Formica vagans Jerdon, 1851. Madras Jour. Lit. Sci. 17: 124. ♂.
Formica gracilescens Nylander, 1856. Ann. Sci. Nat. Zool. 5: 73. ♀.
Paratrechina currens Motschulsky, 1863. See. Nat. Moscou, Bui. 36: 14. ♂.

Taxonomy: Bingham, 1903. Fauna of British India 2: 326-327 (each caste). —Smith, 1936.
Puerto Rico Univ., Jour. Agr. 20: 865, 869-870. —Wheeler and Wheeler 1953. Ent. Soc.
Amer., Ann. 46: 143 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 87
(Polynesia).

Biology: Marlatt, 1928. U. S. Dept. Agr. Farmers' Bui. 740: 6-7. —Phillips, 1934. (Hawaii
Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bui. 15: 18-19 —Turner, 1940. Conn. Agr.
Expt. Sta. Bui. 434: 311-312. —Fox and Garcia-MoU, 1961. Jour. Econ. Ent. 54: 1065-1066.
—Brown, 1964. Ent. News 75: 14-15. —Smith, 1965. U. S. Dept. Agr. Tech. Bui. 1326, pp.
74-76 (economic importance).

Genus PARATRECHINA Subgenus NYLANDERIA Emery

Nylanderia Emery, 1906. Soc. Ent. de Belg., Anrt. 50: 134.

Type-species: Formica vividula Nylander. Orig. desig.

Members of this subgenus are in need of study. Existing keys are not adequate for species
identification, and a number of the names may be wrongly applied. Species determination has al-
ways been considered difficult and male genitalia are believed to offer help in separating spe-
cies. The segregates listed here are essentially as those proposed by Creighton, 1950. This sub-
Genus is much better represented in the tropical regions of the world, and there are many tramp
species which make the taxonomy more confusing.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool, Bui. 104: 402-410.
Biology: Smith, 1965. U. S. Dept. Agr. Tech. Bui. 1326, pp. 76-77 (as house pests).
bourbonica (Forel). S. C, Fla.; Pantropical. Introduced. Probably native to tropical Asia and
spread by commerce to Indian and Pacific Oceans and to New World tropics.
Prenolejris nodifera bourbonica Forel, 1886. Soc. Ent. de Belg., Ann. 30: 210. $, 9, 6.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104: 405 (a var. of

bourbonica in U. S. ?). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 87-89 (Polynesia).

Biology: Smith, 1930. Fla. Ent. 14: 23-24. —Wheeler, 1932. N. Y. Ent. Soc, Jour. 40: 16.
bruesii (AVheeler). Fla., Ala. w. to Okla, Tex., Ariz. (?).

Prenolepis bruesii Wheeler, 1903. Psyche 10: 106. 5, 9, 6.

Taxonomy: Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 211 (larva; Okla.; as
bruesi).

Biology: Smith, 1924. Ent. News 35: 122.
fulva (Mayr). S. Tex. s. to Argentina. Sporadically in greenhouses as far north as N. J.

Probably introduced. References to P. fulva pubens (Forel) from the U. S. pertain to
this species.
Prenolepis fulva Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 698. ♂, ♀.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636-637. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bui. 104: 406-407.

Biology: Marlatt, 1922. U. S. Dept. Agr. Farmers' Bui. 740: 8-9.
melanderi arenivaga (Wheeler). N. J. s. to Fla. w. to Iowa, Tex., Calif.

Prenolepis arenivaga Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 391. 9, Prenolepis {Nylanderia) arenivaga vsir. faisonensis Forel, 1922. Rev. Suisse de Zool. 30:
98. ♂.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 295. —Harper, 1965. Calif. Dept. Agr.
Bui., Ann. Rpt. 45, 54: 81 (Calif, records).

Biology: Smith, 1928. Ent. News 39: 278.




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melanderi melanderi (Wheeler). Tenn. w. to Kans., Tex., Ariz.; Mexico.
Prenolepis melanderi Wheeler, 1903. Psyche 10: 104. 5 , 9 , 6.

Taxonomy: Emery, 1906. Soc. Ent. de Belg., Ann. 50: 132. —Wheeler and Wheeler, 1953. Ent.
Soc. Amer., Ann. 46: 144 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 74.
parvula (Mayr). Mass. s. to Fla. w. to N. Dak., Nebr., Kans., Tex., Ariz., Utah (?). Ecology:
Nests under moss, in logs and stumps, beneath stones, or in open grassy areas where
the nest may be surmounted by a small crater.
Prenolepis parvula Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 947. 5, 9, 3.
Prenolepis (Nylanderia) parvula var. grandula Forel, 1922. Rev. Suisse des Sci. Nat., Bui.
20:348. ♀.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636 (worker, male). — Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 100. — Buren, 1944. Iowa State Col., Jour. Sci. 18:
295. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 211 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 390. —Talbot, 1934. Ecology 15:
420-422. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 295, 306. —Cole, 1940. Amer. Midland
Nat. 24: 66. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 175-176.
vividula guatemalensis (Forel). S. Ariz.; Central Amer. Possibly introduced. According to

Snelling (correspondence), the Ariz, record for guatemalensis is based on misidentified
areniva^a.
Prenolepis vividula vividula var. guatemalensis Forel, 1884. Soc. Vaud. des ScL Nat., Bui.
20:348. ♀.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. ffist., Bui. 21: 392 (worker, male).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 342. —Cole, 1934. Ent. Soc. Amer.,
Ann. 27: 401.
vividula vividula (Nylander). Fla., Miss. Sporadically in greenhouses in northern areas as far

as Canada. Introduced.
Formica vividula Nylander, 1846. Acta Soc. Fenn. 2: 900. 5, 9, Formica perminuta Buckley, 1866. Ent. Soc. Phila., Proc. 6: 162. 5 . Synonymy

questionable.
Formica picea Buckley, 1866. Ent. Soc. Phila., Proc. 6: 163. 5 . Synonymy questionable.
Formica (Tapinoma) terricola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 168. 5, 9, 6.

Synonymy questionable.

Taxonomy: Emery, 1906. Soc. Ent. de Belg., Ann. 50: 130 (each caste). —Emery, 1910. Deut.
Ent. Ztschr., p. 131 (each caste).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 74. — Marlatt, 1922. U. S. Dept.

Agr. Farmers' Bui. 740: 7.

Genus PRENOLEPIS Mayr

Prenolepis Mayr, 1861. Die Europaischen Formiciden, pp. 26, 52.

Type-species: Tapinoma nitens Mayr. Desig. by Bingham, 1903.

These ants usuaUy nest in the soil in exposed situations or under cover in small to moderate
sized colonies. Workers feed on honeydew, secretions of floral and extrafloral nectaries, exu-
dates from galls, earthworms, arthropods, and ripened or decaying fruits. The ants often invade
houses in search of food, and they sometimes forage even at freezing temperatures. Repletes
are conmion. Males and females overwinter in the nest and are among the first of ants to take
their nuptial flights in the spring.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 635-637. —Wheeler, 1930. Ent. Soc.
Amer., Ann. 23: 1-26.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 142 (larvae).

Prenolepis imparls
***authority mismatch
arlzonlca Wheeler. S. Ariz.

Prenolepis imparts var. aiizonica Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 22. 5, 9, cJ.




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imparis califomica Wheeler. Oreg., Nev., Calif. Ecology: Reported to feed on succulent tissue
of fruit and on fruit buds, damaging these by eating out the centers and leaving only
the scales.
Prenolepis imparis var. califomica Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 23. 5, 9, 6.

Taxonomy: Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Bui. 342: 27.

Biology: Essig, 1926. Ins. West. No. Amer., p. 866. —Mallis, 1941. South. Calif. Acad. Sci.,
Bui. 40: 77-78.

Prenolepis imparls
***authority mismatch
coloradensis Wheeler. Colo., N. Mex.

Prenolepis imparis var. coloradensis Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 22. ♂.

Biology: Gregg, 1963. Ants of Colo., pp. 496-498.
imparis imparis (Say). Conn., Ont. s. to Fla. w. to Wis., Iowa, Mo., Okla., Tex., N. Mex., Ariz.

An annoying house infesting ant with habits similar to those given for the genus.
Formica imparis Say, 1836. Boston Jour. Nat. Hist. 1: 287. ♀, ♂.

Tapinoma polita Smith, 1855. Ent. Soc. London, Trans. 3: 112. 9. Synonymy questionable.
Formica {Tapinoma) Wichita Buckley, 1866. Ent. Soc. Phila., Proc. 6: 169. 5. Synonymy

questionable.
Prenolepis nitens var. americana Forel, 1891. In Grandidier, Hist. Madagascar, v. 20, p.

94. ♂.
Prenolepis imparis var. minuta Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636. 5, cJ.
Prenolepis imparis var. testacea Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636. ♀, ♂.
Prenolepis imparis var. pumila Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 21. 5, cJ.

Taxonomy: Wesson and Wesson, 1940. Amer. Midland Nat. 24: 100. —Cole, 1940. Amer.
Midland Nat. 24: 66, 67. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 142-143
(larva).

Biology: Smith, 1924. Ent. News 35: 122. —Smith, 1928. Ent. News 39: 278. —Dennis, 1941.
Ent. Soc. Amer., Ann. 34: 82-86. —Talbot, 1943. Ecology 24: 31-44 (population studies).
—Talbot, 1943. Ecology 24: 345-352. (response to temperature and humidity changes).
— Tarpley, 1965. N. Y. Ent. Soc., Jour. 73: 6 (nuptial flight). —Smith, 1965. U. S. Dept.
Agr., Tech. Bui. 1326, pp. 78-79 (economic importance).

Genus MYRMECOCYSTUS Wesmael

Myrmecocystiis Wesmael, 1838. Brussels Acad. Roy. de Belg., Bui. de CL des Sci. 5: 769.
Type-sjjecies: Myrmecocystzis mexicanus Wesmael. Monotypic.

This genus is native to Mexico and western United States and is especially typical of the arid
plains and deserts. The ants nest in soil, the nest being surmoimted by a small crater, usually in
medium-sized colonies of 1,000 to 3,000 workers. Some forms are apparently entirely predaceous
or carnivorous; others live on honeydew and nectar obtained from plants or the secretion of
galls. They may be diurnal or nocturnal in their activity. Repletes are known to occur in many
forms. Ants of this genus are commonly called "honey ants" because of the honeylike substance
stored in the gasters of the repletes. Fluid stored in repletes is largely for use by adults of the
colony with little or none for the larvae. Larval food consists of dead insects brought in by
foraging workers. Owing to the high degree of polymorphism in many forms, these ants are not
easily determined specifically without large series of workers, especially major ones.

Since this section was completed, Snelling (1976) published a revision of Myrrmcocystus, the
results of which cannot be entirely incorporated into this catalog at this date. Snelling recognizes
27 species in three subgenera, the typical subgenus, subgenus Endiodioctes Snelling
(type-species: Myrmecocystus melliger Forel), and subgenus Eremnocystus Snelling
(type-species: Myrmecocystus creightoni Snelling). Except for their arrangement by subgenus,
all species treated by Snelling that fall within the scope of this catalog are given.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. SystenrL 7: 666-667. —Wheeler, 1908. Amer. Mus.
Nat. Hist., Bui. 24: 345-397. —Wheeler, 1912. Psyche 19: 172-181. —Snelling, 1976. Nat.
Hist. Mus. of Los Angeles Co., Sci. Bui. 24: 1-163 (also biology).

Taxonomy: Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 1-9 (melliger complex).
—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larvae).



1446 Hymenoptera in America Nori;h of Mexico

Biology: Snelling, 1968. Los Angeles Co. Mus., Natur. Hist. Quart. 7: 14-18.

Myrmecocystus colei Snelling
S. Calif.

Myrmecocystus (Eremnocystus) colei Snelling, 1976. Nat. Hist. Mus. of Los Angeles Co.,
Sci. Bui. 24: 94-97. 5, 9,

Myrmecocystus creightoni Snelling
S. Calif.

Myrmecocystus creightoni Snelling, 1971. Los Angeles Co. Mus., Contrib. Sci. 214: 6-11. ♀.
9,6.

Taxonomy: Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva; misidentified as
M. ItLgubris).

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 120.

Myrmecocystus depilis Forel
W. Tex., N. Mex., Ariz., s. Nev.; Mexico. Confused with mimicus in the earlier
literature.
Myrmecocystus melliger var. depilis Forel, 1901. Soc. Ent. de Belg., Ann. 45: 135. ♂.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 354. —Wheeler, 1912. Psyche 19:
173.

Myrmecocystus ewarti Snelling
S. Calif.

Myrmecocystus ewarti Snelling, 1971. Los Angeles Co. Mus., Contrib. Sci. 214: 2-6. ♂, ♀.
S.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 120.

Myrmecocystus flaviceps Wheeler
Utah, Ariz., s. Nev., s. Calif.; Mexico.

Myrmecocystus yuma wsr. flaviceps Wheeler, 1912. Psyche 19: 174, 177. ♂.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 121-122.

Myrmecocystus hammettensis Cole
Idaho, Nev., e. central Calif.

Myrmecocystus hammettensis Cole, 1938. Amer. Midland Nat. 19: 678. 5, 9, d.

Myrmecocystus kathjuli Snelling
S. Calif.

Myrmecocystus {Endiodioctes) kathjuli SneUing, 1976. Nat. Hist. Mus. of Los Angeles Co.,
Sci. Bui. 24: 59-62. 5, 9, cJ.
kennedyi Cole. S. Idaho, s.w. Utah, Ariz., s.e. Oreg., Nev., Calif.; Mexico. This species was
called M. semirufus in the Uterature prior to Snelling, 1969.
Myrmecocystus melliger semirufus var. kennedyi Cole, 1936. Ent. News 47: 119. 5, 9, 6.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 355, 368-369 (semirufus; also
biology). —Wheeler, 1912. Psyche 19: 174, 176 (semirufus; also biology). —Cole, 1938.
Amer. Midland Nat. 20: 371, 372. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61:
213 (larva; as semirufus). — Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 6.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 20 (semirufus). — Wheeler and
Wheeler, 1973. Ants of Deep Canyon, p. 122.

Myrmecocystus koso Snelling
S. Nev., s. Calif.

Myrmecocystus (Endiodioctes) koso Snelling, 1976. Nat. Hist. Mus. of Los Angeles Co.,
Sci. Bui. 24: 74-78. 5, 9, d.
lug^ubris Wheeler. S. Nev., s. Calif.

Myrmecocystus lugubris Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 98. ♂.

Taxonomy: Wheeler, 1912. Psyche 19: 174, 176. — Creighton, 1956. Amer. Mus. Novitates
1807: 1.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 21.

Myrmecocystus melliger
***authority mismatch
ForeL W. Tex.; Mexico. Used as food and medicine by Indians.

Formica melligera Llave, 1832. Reg. Trim. o. Collect. Mem. Hist. Lit., p. 463. ♀.

Questionable placement.
Myrmecocystus melliger Forel, 1886. Soc. Ent. de Belg., Ann. 30: 201. ♂.
Myrmecocystus melliger mendax var. comntus Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.
24:352. 5, 9, 6.



Superfamily FORMICOIDEA 1447

Taxonomy: Wheeler, 1912. Psyche 19: 173, 175. — Snelling, 1969. Los Angeles Co. Mus.,
Contrib. Sci. 170: 4. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 211-213
(larva).

Myrmecocystus mendax Wheeler
Tex., Colo., N. Mex., Ariz., s. Nev., s. Calif.; Mexico.

Myrmecocystus melliger mendax Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 351. ♂, ♀.

6.
Myrmecocystus melliger orbiceps Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 349. ♂, ♀.

Taxonomy: Wheeler, 1912. Psyche 19: 173. —Cole, 1954. Tenn. Acad. Sci., Jour. 29: 284.
—Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 2.

Biology: Wheeler, 1910. Ants, p. 376. —Cole, 1942. Amer. Midland Nat. 28: 386. —Gregg,
1963. Ants of Colo., pp. 643-645, 645-648 (misident. as comatus, p. 643-645).

Myrmecocystus mexicanus Wesmael
W. Tex., Colo., N. Mex., Utah, Ariz., Nev., Calif.; Mexico.

Myrmecocystus mexicanus Wesmael, 1838. Brussels Acad. Roy. de Belg., Bui. CI. des Sci.

5: 770. ♂.
Myrmecocystus Melliger var. hortus-deorum. McCook, 1881. Acad. Nat. Sci. Phila., Proc.,
pp. 65, 69. 5, replete 5, 9, <5.

Taxonomy: Forel, 1886. Soc. Ent. de Belg., Ann. 30: 202. —Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 666. —Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 356-360. —Wheeler, 1912.
Psyche 19: 173, 178. —Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva).

Biology: McCook, 1881. Acad. Nat. Sci. Phila., Proc., pp. 17-77. —McCook, 1882. The Honey
Ants of the Garden of the Gods and the Occident Ants of the American Plains, pp. 17-74.
— Mallis, 1941. South. Calif. Acad. Sci., Bid. 40: 81. —Gregg, 1963. Ants of Colo., pp.
648-651. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 122-124.

Myrmecocystus mimicus Wheeler
S.W. Kans., w. Okla., w. Tex., N. Mex., Ariz., Calif.; Mexico.

Myrmecocystus melliger mim.icus Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 353. ♂.

9,d.
Myrmecocystus melliger mAmicus var. jesuita Wheeler, 1908. Amer. Mus. Nat. Hist., Bui.

24: 354. ♂.
Myrmecocystus melliger lomaensis Wheeler, 1912. Psyche 19: 174. 5, replete 5,9.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104: 446. —Wheeler and
Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva).

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27: 401^02. —Mallis, 1941. South. Calif. Acad. Sci.,
Bui. 40: 80. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 22 (Nev. Test Site).
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 124-125. — Holldobler, 1976.
Science 192: 192-194 (tournaments and slavery).

Myrmecocystus navajo Wheeler
Colo., w. Tex., N. Mex., Utah, Ariz., s. Nev., s. Calif.

Myrmecocystus mexicanus navajo Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 360. ♂.
9.

Taxonomy: Wheeler, 1912. Psyche 19: 173, 179.

Biology: Cole, 1942. Amer. Midland Nat. 28: 386. —Gregg, 1963. Ants of Colo., pp. 651-653.
— Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 125.

Myrmecocystus placodops
***authority mismatch
ForeL Tex. N. Mex., Ariz.; Mexico.

Myrmecocystus melliger var. placodops Forel, 1908. Soc. Vaud. des Sci. Nat., BuL 44: 70.

5.

Taxonomy: Wheeler, 1912. Psyche 19: 173. —Snelling, 1969. Los Angeles Co. Mus., Contrib.
Sci. 170: 6.

Myrmecocystus pyramicus Smith
Idaho, s.e. Oreg., Nev.

Myrmecocystus pyramicus Smith, 1951. Great Basin Nat. 11: 91-94. ♂.

Taxonomy: Cole, 1957. N. Y. Ent. Soc., Jour. 65: 129-130 (female, male).

Myrmecocystus romainei Cole
W. Kans., w. Okla., w. Tex., Colo., N. Mex., Utah, Ariz., Nev.; Mexico.
Myrmecocystus melliger semirufus var. romainei Cole, 1936. Ent. News 47: 120. ♂.




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Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 285 (as semirufa, in part). —Gregg, 1963. Ants
of Colo., pp. 653-655 (as semirufus, in part).

Myrmecocystus semirufus Emery
S. Calif.; Mexico.

Myrmecocystus melliger var. semirufus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 667.

5.

Taxonomy: Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 4-6 (The species

previously considered to be M. semirufus by Wheeler and subsequent authors must now
be known as M. kennedyi Cole.).

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 125-126 (misident. as

Myrmecocystus placodops Forel
).

Myrmecocystus tenuinodis Snelling
S. Nev., s.w. Ariz., s. Calif.; Mexico.

Myrmecocystus {Eremnocystus) tenuinodis Snelling, 1976. Nat. Hist. Mus. of Los Angeles
Co., Sci. Bui. 24: 107-110. ♂, ♀.

Myrmecocystus testaceus Emery
Idaho, Utah, Wash. Oreg., Nev., Calif.; Mexico.

Myrnnecocystus melliger var. testaceus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 667.

5.
Myrmecocystus m^xicanus mojave Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 360. ♂.
Myrmecocystus m^xicanus idahoensis Cole, 1936. Ent. News 47: 118. 5, 9, 6.

Taxonomy: Wheeler, 1912. Psyche 19: 173, 179-181. —Snelling, 1969. Los Angeles Co. Mus.,
Contrib. Sci. 170: 6. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva).

Biology: Wheeler, 1910. Ants, p. 349. —Leonard, 1911. San Diego Soc. Nat. Hist., Trans. 1:
91-92. —Cole, 1934. Ent. Soc. Amer., Ann. 27: 403. — Mallis, 1941. South. Calif. Acad. Sci.,
Bui. 40: 81. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 22 (Nev. Test Site).
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 126-127.

Myrmecocystus wheeleri Snelling
S. Calif.

Myrmecocystus wheeleri Snelling, 1971. Los Angeles Co. Mus., Contrib. Sci. 214: 11-15. ♀.
9,6.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 127.

Myrmecocystus yuma Wheeler
W. Ariz., s. Nev., s. Calif.; Mexico.

Myrmecocystus yuma Wheeler, 1912. Psyche 19: 174, 176. ♂.

Taxonomy: Creighton, 1956. Amer. Mus. Novitates 1087: 1.

Tribe FORMICINI
Genus FORMICA Linnaeus

Formica Linnaeus, 1758. Syst. Nat., Ed. 10, v. 1, p. 579.

Type-species: Formica rufa Linnaeus. Desig. by Curtis, 1839.
Serviformica Forel, 1913. Soc. Ent. de Belg., Ann. 57: 361.

Type-species: Formica fusca Linnaeus. Orig. desig.
Raptifarmica Forel, 1913. Soc. Ent. de Belg., Ann. 57: 361.

Type-species: Formica sanguinea Latreille. Orig. desig.
Formica subg. Neoformica Wheeler, 1913. N. Y. Acad. Sci., Ann. 23: 82.

Type-species: Formica pallide-fulva Latreille. Desig. by Wheeler, 1913.
Coptoformica Mueller, 1923. Soc. Adriat. di Sci. Nat. Trieste, Bol. 28: 133.

Type-species: Formica exsecta Nylander. Desig. by Donisthorpe, 1941.
Adformica Lomnicki, 1925. Polskie Pismo Ent. 3: 164.

Type-species: Formica exsecta Nylander. Desig. by Donisthorpe, 1941.

This is a large and complex holarctic genus, and the largest genus of ants in America north of
Mexico. Various habit patterns are found in this genus including slave-making, temporary social
parasitism of various types, and various methods of nest founding and nest construction. Sub-
genera are not recognized in the following listing, only species groups which more or less reflect
the groupings by Creighton (1950) and Buren (1968). Proformica, apparently a valid genus found
in Europe, is deleted from the North American fauna, and those species previously listed in that




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subgenus along with a number of species excluded from Raptiformica by Buren are here listed
in the species group tieogagates. Those species included in the subgenus Raptiformica by Buren
(1968) are listed here in the species group sanguinea. The species previously included in the
subgenus Neoformica are listed here in the pallidefulva species group. The other species
groups, microgyna, exsecta, fusca, and rufa, are essentially the same as before. Further studies
may clarify the species groupings and determine the necessity of establishing subgenera.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643-654, 657-665. —Wheeler, 1913.
Harvard Univ., Mus. Comp. Zool., Bui. 53: 387-399, 401-536, 560-565. — Creighton, 1940.
Amer. Mus. Novitates 1055: 1-10 (rufa subspp.). —Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bui. 104:450-552. — Francoeur, 1973. Ent. Soc. du Quebec, Mem. 3, 316 pp.
(fusca group).

Taxonomy: Yarrow, 1950. Intematl. Comm. Zool. NomencL, Bui. Zool. Nomencl. 4: 408.
—Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 160 (larvae). —Yarrow, 1955.
Intematl. Comm. Zool. Nomencl., Bui. Zool. Nomencl. 9: 313-318 (type-species). —Wilson
and Brown, 1955. Psyche 62: 108-129 (notes on sanguinea and neogagates groups).
—Gregg, 1964. Colo. Univ. Mus., Leaflet No. 13, pp. 59-69 (distribution in mountains of
Colo.). —Gregg, 1969. Ent. Soc. Wash., Proc. 71: 38-49 (geographical distribution of genus
in world). —Buren, 1968. Ga. Ent. Soc., Jour. 3: 25-40 (fundamental taxonomic problems;
subg. Raptiformica).

Biology: (iosswald, 1957. Insectes Sociaux 4: 335-348 (caste determination). — Kannowski,
1959. Insectes Sociaux 6: 147. —Kannowski and Johnson, 1969. Anim. Behaviour 17:
425:429 (male patrolling behavior and sex attraction). — Rosengren, 1971. Acta Zool.
Fennica 133, 106 pp. (route fidelity, visual memory, recruitment behavior).

Morphology: Osman and Kloft, 1961. Insectes Sociaux 8: 383-395 (venom).

Species Group Neogagates

Species of this group nest in small colonies in the soil, sometimes under stones or beneath
other objects. They are commonly enslaved by members of the microgyna and sanguinea
groups.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 108-129. —Buren, 1968. Ga. Ent. Soc., Jour.
3: 25-40.

Formica bradleyi Wheeler
Man., Minn., Iowa w. to Alta., Mont., Wyo., Colo. Ecology: Nests only in
sand or sandy soil among grass cltunps or with low irregular mounds.
Formica bradleyi Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 389, 423. 9, Formica (Proformica) neogagates neogagates var. morbida Wheeler, 1913. Harvard Univ.,
Mus. Comp. Zool., Bui. 53: 400, 538. 9, 9.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 535. —Wheeler and Wheeler,
1953. Ent. Soc. Amer., Ann. 46: 168-169 (larva). — WUson and Brovm, 1955. Psyche 62:
126-127.

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 258-259. — Beamer and

Michner, 1950. Kans. Ent. Soc., Jour. 23: 110-113 (relationship with leafhoppers).

—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 206-209. —Gregg, 1963. Ants of Colo.,

pp. 602-606. — Snelling, 1969. Ent. Soc. Wash., Proc. 71: 196-197. — Halverson, et al. 1976.

Kans. Ent. Soc., Jour. 49: 280-303 (natural history of the sandhill ant).

Formica lasioides Emery
N. S., Que. w. to B. C. s. to Mass., Mich., S. Dak., Colo., N. Mex., Ariz., Calif.
Ecology: Commonly foimd in grasslands where it nests under stones or in nests vdth
exposed entrances or small craters, but also found in other habitats.

Formica lasiodes Emery, 1893. Zool. Jahrb., Abt. f. System 7: 646, 664. ♂.

Formica lasoides var. picea Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 335. ♂. Preocc.
by Nylander, 1846.

Formica lasoides var. vetula Wheeler, 1912. Psyche 19: 90. N. name for picea Emery.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 400, 540-541 (each
caste). —Gregg, 1944. Ent. Soc. Amer., Ann. 37: 472. —Buren, 1944. Iowa State Col., Jour.
Sci. 18: 299.




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Biology: Cole, 1942. Amer. Midland Nat. 28: 384. —Wheeler and Wheeler, 1944. N. Dak. Hist.
Quart. 11: 268-269. — Kincaid, 1963. Amer. Micros. Soc, Trans. 82: 101-105 (as pollinators of
plants). —Wheeler and Wheeler, 1963. Ants of N. Dak. pp. 209-211, 502. —Gregg, 1963.
Ants of Colo., pp. 498-500, 502. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7:

23 (Nev. Test Site).

Formica limata Wheeler
Minn., N. Dak., Colo., N. Mex., Utah, Nev. Ecology: Commonly nests in
grasslands under stones or in crater nests; tolerant of dry situations.
Formica (Proformica) limata, Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53:
400,541. ♂.

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 268. —Gregg, 1946. Amer.
Midland Nat. 35: 750. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 211-212. —Gregg,
1963. Ants of Colo., pp. 502, 504. —Cole, 1%6. Brigham Young Univ., Sci. Bui., Biol. Ser. 7:

24 (Nev. Test Site).

Formica manni Wheeler
Idaho, Utah, Wash., Oreg., Nev., Calif. Ecology: Nests are often under stones
in gravelly or sandy soil of desert areas.
Formica manni Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 389, 420. ♂, ♀.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 127-128.

Biology: Cole, 1942. Amer. Midland Nat. 28: 376, 378-379.

Formica neogagates Emery
N. S., Que. w. to Alaska s. to N. C, 111., Iowa, Nebr., N. Mex., Ariz., Calif.
Ecology: Nests most often in grasslands in dry and stony situations, under stones or in
the open with or without an irregular mound or crater.
Formica fusca subpolita var. neogagates Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646,

661. 5, 9, i.
Formica {Proformica) neogagates neogagates var. vinculans Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool., Bui. 53: 400, 539. ♂, ♀.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 400, 536-538 (each
caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22: 596. —Wheeler
and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 158-159 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 401. —Wesson and Wesson, 1940.
Amer. Midland Nat. 24: 102. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 268.
— Buren, 1944. Iowa State Col., Jour. Sci. 18: 299. —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 212-214. —Gregg, 1963. Ants of Colo., pp. 503-505. —Cole, 1966. Brigham
Young Univ., Sci. Bui., Biol. Ser. 7: 26 (Nev. Test Site).

Formica obtusopilosa Emery
Minn. w. to Alta. s. to Nebr., N. Mex., Utah, Nev. Ecology: Small colonies

are found under stones or in exposed soil usually with irregular moimds or craters in

meadows and grasslands.
Formica sanguinea obtusopilosa Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 648. ♂.
Formica munda Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 267. ♂, ♀.
Formica munda var. alticola Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 534. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 389, 414-418. — WUson
and Brown, 1955. Psyche 62: 128.

Biology: Cole, 1942. Amer. Midland Nat. 28: 378. —Wheeler and Wheeler, 1963. Ants of N.
Dak, pp. 217-219. —Gregg, 1963. Ants of Colo., pp. 608-611. —Cole, 1966. Brigham Young
Univ., Sci. Bui., Biol. Ser. 7: 26 (Nev. Test Site).

Formica oregonensis Cole
Oreg. (Pendleton).

Formica oregonensis Cole, 1938. Amer. Midland Nat. 20: 368. ♂.

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31: 212-214.

Biology: Kincaid, 1963. Amer. Micros. Soc., Trans. 82: 101-105 (pollinators of plants).

Formica perpilosa Wheeler
Wyo., Colo., Kans., Okla., Tex. w. to Calif.; Mexico. Ecology: Usually
constructs crater nests in the soil in grasslands and open fields.
Formica fusca subpolita var. perpilosa Wheeler, 1902. Soc. Cient. "Antonio Alzate" Mem.
y Rev. 17: 141. ♂, ♀.




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Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 389, 421-423 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376-378. —Wilson and Brown, 1955. Psyche
62: 128.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 534. —Cole, 1934. Ent. Soc.
Amer., Ann. 27: 401. — Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 85. — LaBerge, 1952.
Kans. Ent. Soc, Jour. 25: 59. —Gregg, 1963. Ants of Colo., pp. 611-613.

Species Group Paludefulva

Members of this group nest in the soil under stones or at the base of tufts of grass. There is
usually no mound or crater marking the nest. They serve as hosts of slave-making species of
Formica.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104: 543-552 (subg.
Neoformica).

Formica archboldi Smith
Ga., Ala., Fla.

Formica palHde-fulva archboldi Smith, 1944. Fla. Ent. 27: 16. 2.

Biology: Schneirla, 1944. Amer. Mus. Novitates 1261: 1-2.

Formica pallidefulva
***authority mismatch
nitidiventris Emery. Ont., Que. s. to Ga. w. to Wis., S. Dak, Wyo., Colo., N. Mex.
Formica pallide-fulva nitidiventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 656.

5, ?, 6.
Formica pallide-fulva schaufussi war. incerta Emery, 1893. Zool. Jahrb., Abt. f. System. 7:

645,655. 5, 9, 6.
Formica pallide-fulva fuscata Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 656. 2.
Formica pallide-fulva delicata Cole, 1938. Amer. Midland Nat. 20: 369. ♂, ♀.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 401. —Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool., Bui. 53: 401, 554-557. —Wheeler, 1917. Conn. State Geol. and Nat.
Hist. Survey Bui. 22: 595, 598. — Buren, 1944. Iowa State Col, Jour. Sci. 18: 299, 309-310.
—Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 159-160 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 370-371. —Wheeler, 1917. Ind. Acad.
Sci., Proc. 26: 465. — Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 211-212. —Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 102. —Talbot, 1946. Ecology 27: 65-70. — Schread
and Chapman, 1948. Conn. (State) Agr. Expt. Sta. Bui. 515: 18. —Talbot, 1948. Ecology 29:
316-325. —Gregg, 1963. Ants of Colo., pp. 628-630.

Formica pallidefulva
***authority mismatch
pallidefulva Latreille. N. Y., N. J. s. to Fla. w. to Colo., Okla., Tex.
Formica pallide-fulva Latreille, 1802. Hist. Nat. Fourmis, p. 174. ♂.
Formica pallide-fulva var. su^cinea Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 369. ♂.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 656-657. —Wheeler, 1913.
Harvard Univ., Mus. Comp. Zool., Bui. 53: 401, 548-552.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 298, 306. —Cole, 1940. Amer. Midland Nat.
24: 81. —Schneirla, 1944. Amer Mus. Novitates 1261: 2-3. —Gregg, 1963. Ants of Colo., pp.
625-628.

Formica schaufussi
***authority mismatch
dolosa Wheeler. Va. s. to Fla. w. to Iowa, Colo., Tex.

Formica pallide-fulva schaufussi var. meridionalis Wheeler, 1904. Amer. Mus. Nat. Hist.,

Bui. 20: 370. ♂. Preocc. by Ruzsky, 1889.
Formica pallide-fulva schaufussi var. dolosa Wheeler, 1912. Psyche 19: 90. N. name for

Formica meridionalis Wheeler


Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 401, 554 (worker,
female). —Cole, 1940. Amer. Midland Nat. 24: 73, 79. —Buren, 1944. Iowa State Col., Jour.
Sci. 18: 299.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 299, 306. —Gregg, 1963. Ants of Colo., pp.
630-632.

Formica schaufussi
***authority mismatch
schaufussi Mayr. Ont. s. to N. C, Tenn. w. to Wis., Iowa.

Formica Schaufussi Mayr, 1886. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 53: 493. ♀.




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Taxonomy: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 370. —Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool, Bui. 53: 401, 552-553 (each caste). —Cole, 1940. Amer. Midland
Nat. 24: 73.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 400. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bui. 22: 598. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 298,
306.

Species Group Fusca

Members of this group nest in the soU, and the nest is commonly started under objects or at
the base of tufts of grass. Excavated soil may be spread out in the form of an irregular crater or
low mound about the nest openings. Because of their ubiquity and usual docility, they are com-
mon hosts for many of the slave-making species of Formica.

Revision: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, 316 pp.

Formica accreta Francoeur
B. C, Mont., Idaho, Wash., Oreg., Calif.

Formica accreta Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 182-189. 5 , 9 , 3.

Formica aerata Francoeur
Oreg., Nev., Calif. Ecology: Found in sandy soil where it nests under rocks.

Formica aerata Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 116-122. ♂, ♀.

Formica altipetens Wheeler
Mont., Wyo., Colo., N. Mex., Idaho, Utah, Ariz., Calif. Ecology: Nests in
forested and open areas in the soil; low moimds are sometimes built, but nests are also
under objects.
Formica cinerea cinerea var. altipetens Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bui. 53: 399, 523. 5, 9,
Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 52-60.

Biology: Gregg, 1963. Ants of Colo., pp. 507-509. —Funk, 1975. Lepidop. Soc., Jour. 29:
261-262 (association with Lycaenidae).

Formica argentea Wheeler
Que. w. to B. C. s. to S. C, Ohio, 111., Iowa, S. Dak., N. Mex., Ariz., Calif.

Ecology: Found in ojjen or semi-open situations usually in sandy soil under rocks or

with a low mound.
Formica fusca var. argentata Wheeler, 1902. Amer. Nat. 36: 952. ♂. Preocc. by Fabricius,

1804.
Formica fusca var. argentea Wheeler, 1912. Psyche 19: 90. N. name for argentata

Wheeler.
Form,ica fusca var. blanda Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 398,

510. ♀.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 141-152 (also ecology).

Biology: Gregg, 1963. Ants of Colo., pp. 521-525.

Formica canadensis Santschi
Man., Sask., N. Dak., S. Dak., Kans., Alta., Wyo., Colo., N. Mex., Idaho,

Utah, Ariz., Calif. Ecology: Nests in open or wooded areas in the soil, sometimes with a
low mound. This species has been confused with lepida in much of the literature.
Formica cinerea var. canadensis Santschi, 1913. Soc. Ent. de Belg., Ann. 57: 435. ♂, ♀.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 60-67.

Formica foreliana Wheeler
S. Ariz.

Formica fareliana Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., BuL 53: 391, 451. ♂.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 234-238.

Formica fusca Linnaeus
Newfoundland (insular) w. to Yukon s. to N. C, 111., Iowa, S. Dak., N. Mex.,

Ariz., Calif.; Holarctic. Ecology: Nests in a wide variety of situations, in forests or open
areas, under rocks, logs, in soil, or in rotting wood. Two forms are recognized by
Francoeur (1973), form subaenescens which is mostly eastern and form mardda which
is mostly western in distribution.

Formica fusca Linnaeus, 1758. Syst. Nat., Ed. 10, v. 1, p. 580.

Formica fusca var. subaenescens Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646,
659-660. $, 9.

Formica fusca fusca var. mardda Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui.
53:398,503. ♂, ♀.




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Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104: 532. — Wheeler and
Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 164-165 (larva). —Yarrow, 1954. Soc. Brit. Ent.,
Trans. 11: 229-244. — Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 189-201.

Biology: King, 1949. Iowa Acad. Sci., Proc. 56: 367-370 (in mixed colony with Formica reflexa
Buren). —King, 1951. Iowa Acad. Sci., Proc. 58: 487-489. — Kannowski, 1959. Insectes
Sociaux 6: 141-142. — Wallis, 1960. Insectes Sociaux 7: 187-190 (spinning movements of
larvae). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 264-268. —Gregg, 1963. Ants of
Colo., pp. 514-525. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 23 (Nev. Test
Site).

Formica glacialis Wheeler
Newfoundland (insular) s. to N. Y. w. to Man., N. Dak., 111. Ecology: Found
in woods or open fields where it builds its nests in the soil and with a low mound
commonly covered with grass. This species has been confused with fusca L. in much of
the literature.
Formica fusca var. glacialis Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 624. 5, 9, 6.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 151-161 (also ecology).

Formica gnava Buckley
Tex., Colo., N. Mex., Utah, Ariz., Nev., Calif.; Mexico. Ecology: Found in desert
and semi-desert areas or oi)en woods where it nests in the soil, normally under rocks.
Formica gnava Buckley, 1866. Ent. Soc. Phila., Proc. 6: 156. 5, 9, 6.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53: 390, 518-521 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 377, 380, 383. —Francoeur, 1973. Soc. Ent. du
Quebec, Mem. no. 3, pp. 238-245.

Biology: Wheeler, 1902. Tex. Acad. Sci., Trans. 4: 20. —Wheeler, 1917. Amer. Acad. Arts and
Sci., Proc. 52: 541, 550. —Cole, 1934. Ent. Soc. Amer., Ann. 27: 388. — Mallis, 1941. South.
Calif. Acad. Sci., Bui. 40: 86. —Gregg, 1963. Ants of Colo., pp. 539-541.

Formica hewitti Wheeler
Que., Maine w. to B. C. s. to Minn., N. Mex., Utah, Calif. Ecology: Found in
open or semi-open woods and forests where it nests in the soil under rocks or in rotting
wood.
Formica hewitti Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 552. 5, 9, <5.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 208-215.

Biology: Gregg, 1963. Ants of Colo., pp. 526-527.

Formica lepida Wheeler
Calif. Has been found only in the north coastal region.

Formica cinerea cinerea var. lepida Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui.
53: 399, 526. ♂.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 122-125.

Formica longipilosa Francoeur
Calif. (Mendocino Co.).

Formica longipilosa Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 125-128. ♂.

Formica microphthalma Francoeur
Calif. Ecology: In mountains; may construct small crater nests in
the soU.
Formica microphthalma Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 201-207. ♀.

a.

Formica moki Wheeler
Utah, Ariz., Nev., Calif. Ecology: Nests in soil under rocks; the nest is

unmarked and the entrance is usually a hole near a bush or tree. Francoeur (1973) chose
to use the name occidua for this species because occidua has been used more in the
literature; however, the case has not been referred to the Commission.

Formica moki Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 343. ♂.

Formica rufibarbis var. occidentalis Wheeler, 1910. Ants, p. 570. Nomen nudum. Preocc.
by Buckley, 1866.

Formica rufibarbis var. occidua Wheeler, 1912. Psyche 19: 90. Nomen nudum. N. name for

Formica occidentalis Wheeler


Formica rufibarbis var. occidun Wheeler, 1913. Harvard Univ., Mus. Comp. ZooL, Bui. 53:
398,517. ♂, ♀.

Taxonomy: Smith, 1939. Ent. Soc. Amer., Ann. 32: 582-583. —Cole, 1943. Amer. Midland Nat.
29: 183. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 24-25 (male, also




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biological notes). — Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 254-259 (as
occidua).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 550. — Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 30. —Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 86.
—Cole, 1942. Amer. Midland Nat. 28: 385. — Wenner, 1959. Amer. Midland Nat. 62: 174.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 128-129.

Formica montana Emery
Ohio, Wis., Ill, Minn., Iowa, Man., N. Dak., S. Dak., Nebr., Kans., Colo.

Ecology: A prairie species; nests in earthen mounds, often in natural hummocks,

commonly covered with grass. Sometimes uses thatching.
Formica fusca subpolita var. ? montana Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646,

663. ♂.
Formica cinerea var. neocinerea Wheeler, 1910. Ants, p. 571. Nomen nudum.
Fonrmica cinerea var. neocinerea Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53:

524. 5, 9, i.
Formica cinerea cinerea var. rutilans Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,

Bui. 53: 399, 525. ♂.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 67-74.

Biology: Wheeler, 1902. Amer. Nat. 36: 948-952. —Wheeler, 1910. Ants, pp. 201, 203, 460-461,
475. —Wheeler, 1917. Psyche 22: 206. — Amstutz, 1943. Ohio Jour. Sci. 43: 172. —Gregg,
1948. Ent. Soc. Wash., Proc. 50: 183-186. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 263-264 (also as altipetens, pp. 260-261). —Gregg, 1963. Ants of Colo., pp. 513-514.

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 455 (chromosome number).

Formica neoclara Emery
N. Dak., S. Dak., Iowa, Kans., Tex. w. and northwest to N. W. T., B. C,

Wash., Oreg., Calif. Ecology: Found in grasslands or open woods where it nests in the
soil, usually preferring sandy soil; nests are sometimes at the base of plants and
sometimes have loose mounds of vegetable debris or excavated soil.
Formica fusca var. neoclara Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646, 661. ♂.
Formica fusca pruinosa Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 548. 5, 9, 6.
Formica fusca pruinosa var. lutescens Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:
549. ♀.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 165 (larva). —Francoeur,
1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 84-94.

Biology: Wheeler, 1910. Ants, pp. 201, 460-461, 463. — LaBerge, 1952. Kans. Ent. Soc., Jour.

25: 59. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 268-271. —Gregg, 1963. Ants of

Colo., pp. 526, 528-530 (also as pruniosa, pp. 537-539).

Formica neorufibarbis Emery
Newfoundland (Labrador) w. to Alaska s. to Mass., Mich., Minn., S. Dak.,
N. Mex., Ariz., Calif. Ecology: A dominant ant in the boreal and alpine forests of North
America; usually nests in rotting wood but occasionaUy in the soU under rocks.

Formica fusca var. neorufibarbis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646, 660. ♂.

Formica fusca var. algida Wheeler, 1915. Psyche 22: 205. ♂, ♀.

Formica fusca fusca var. gelida Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53:
399,505. 5, 9, 6.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 546-547. — Wheeler and
Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 165 flarva). —Francoeur, 1973. Soc. Ent. du
Quebec, Mem. no. 3, pp. 215-228.

Biology: Brown, 1955. Ent. News 66: 47-50. — Kannowski, 1959. Insectes Sociaux 6: 142.

—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 271-273. —Gregg, 1963. Ants of Colo.,

pp. 530-536. —Cole, 1966. Brigham Young Univ., Sci. Bui., BioL Ser. 7: 26 (Nev. Test Site).

Formica occulta Francoeur
Wyo., Colo., N. Mex., Utah, Ariz., Oreg.

Formica occulta Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 94-101. $, 9, 6.
pacifies Francoeur. B. C, Wash., Oreg., Calif.

Formica pacifica Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 131-135. ♂, ♀.




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pilicomis Emery. Calif.; Mexico. Ecology: Found at low elevations in the coast ranges north to
the San Francisco area; makes crater or mound nests in sandy soil, sometimes under
rocks.
Formica pilicomis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646, 664. 5, 9, 6.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 74-80.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27: 401. — Eckert and Mallis, 1937. Calif. Agr.
Expt. Sta. Cir. 342: 7, 29. —Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 85.

Formica podzolica Francoeur
N. S., Que. w. to Alaska s. to Pa., Wis., Iowa, S. Dak., N. Mex., Ariz.,

Calif. Ecology: A species of the boreal and alpine forests of North America. Nests are
in the soil, commonly sandy soil on beaches or shores and are craterUke or moundlike.
Has been confused with F. fusca L. in the literature.
Formica podzolica Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 167-172. 5, 9, S.
sibylla Wheeler. Oreg., Nev., Calif. Ecology: Makes crater nests in the soil, usually in
woodlands.
Formica sibylla Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 399, 530. ♂, ♀.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 263-268.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 551. —Mallis, 1941. South. Calif.
Acad Sci., Bui. 40: 87.

Formica subelongata Francoeur
Calif. (Bumey Junction, Shasta Co.).

Formica subelongata Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 268-270. ♂.

Formica subpolita Mayr
B. C, Idaho, Wash., Oreg., Nev., Calif. Ecology: Usually found in semi-desert

areas where it nests in sandy or gravelly soil; nests may be craterlike or moundlike.
Fcrrmica fusca var. subpolita Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 426. ♂, ♀.
Formica rufiventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 665. ♂. Preocc. by

Fabricius, 1804.
Formica flam,m,iventrls Wheeler, 1912. Psyche 19: 90. N. name for rufiventris Emery.
Formica subpolita var. cam,ponoticeps Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,

Bui. 53: 399, 535. ♂.
Formica subpolita var. ficticia Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53:

561. 5, 9, 6.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 106-115.

Biology: Wheeler, 1910. Ants, pp. 201, 460. —Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.

52: 553-554. —Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 87. —Cole, 1942. Amer.

Midland Nat. 28: 384, 377. —Cole, 1966. Brigham Young Univ., Sci. BuL, Biol. Ser. 7: 26

(Nev. Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 129.

Formica subsericea Say
N. B., Que. s. to Fla. w. to Man., Mont., Iowa, Kans., Mo., Miss. Ecology:
Occurs in open deciduous woodlands where it nests in the soil under stones or leaf
litter; sometimes builds low mounds which may be covered with debris. This species has
been confused with F. fusca L. in the literature.

Formica subsericea Say, 1836. Boston Jour. Nat. Hist. 1: 289. 5, d.

Formica lecontei Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 542. 5, 9, S.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 172-181.
Biology: Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 542. —Wilson and Francoeur,
1974. Fla. Ent. 57: 115-116 (Fla.).

Formica transmontanis Francoeur
Alta., B. C, Mont., Idaho, Calif.

Formica transmontanis Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 135-140. ♂.
9,6.

Formica xerophila Smith
Wash., Utah, Ariz., Calif.

Formica (Neoformica) moki xerophila Smith, 1939. Ent. Soc. Amer., Ann. 32: 583. ♂.
Formica moki grundmanni Cole, 1943. Amer. Midland Nat. 29: 184. ♀.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 259-263.




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SpfeciES Group Exsecta

Ants of this group build large conspicuous mounds in fields, woods, or at the edge of woods.
The nest founding female may behave as a temporary social parasite, but colonies are also
founded by budding.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. ZooL, Bui. 104: 510-515.

Formica exsectoides Forel
N. S., Ont. s. to Ga. w. to Wis., Iowa, Kans., Colo., N. Mex. Ecology: A large

aggregation of mounds of this species may occur in a single locality. Sometimes damages

bark and cambium of small trees and shrubs. Host: Formica fusca L. Often locally

called the Allegheny mound ant.
Formica exsectoides Forel, 1886. Soc. Ent. de Belg., Ann. (C. R.) 30: 38. ♂, ♀.
Formica exsectoides exsectoides var. davisi Wheeler, 1913. Harvard Univ., Mus. Comp.

Zool., Bui. 53: 396, 484. ♂, ♀.
Formica exsectoides exsectoides var. hesperia Wheeler, 1913. Harvard Univ., Mus. Comp.

Zool., Bui. 53: 396, 484. ♀.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 653 (worker, male). —Wheeler,
1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 396, 481-483 (each caste). — Buren, 1944.
Iowa State Col., Jour. Sci. 18: 300. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46:
164 (larva). —Cole, 1954. Tenn. Acad. Sci., Jour. 29: 166.

Biology: McCook, 1877. Amer. Ent. Soc., Trans. 6: 253-295. — Pierson, 1922. Jour. Forestry
20: 325-336. — Manter, 1925. Jour. Econ. Ent. 48: 348-351. —Andrews, 1925. Psyche 32:
75-87 (growth of mounds). —Andrews, 1926. Psyche 33: 127-150. —Andrews, 1929. Ent.
Soc. Amer., Ann. 22: 369-391 (association vdth treehoppers). —Andrews, 1929. Quart. Rev.
Biol. 4: 248-257. —Andrews, 1941. Sci. Monthly 43: 530-533. — Havlland, 1947. Jour. Econ.
Ent. 40: 413-418 (biology and control). — Schread, 1949. Jour. Econ. Ent. 42: 501. —Gregg,
1963. Ants of Colo., pp. 597-599. — Christensen and Quick, 1970. Iowa Acad. Sci., Proc. 77:
207-209.

Morphology: Eisner and Happ, 1964. Science 134: 329-331 (haemorrhage of a coccinellid
beetle and its repellent effect on ants).

Formica opaciventris Emery
N. Dak., Mont., Wyo., Colo., N. Mex. Ecology: May construct either
earthen mounds or thatched nests.
Formica exsectoides var. opaciventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643,
653. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 484-485 (worker, male).
—Gregg, 1952. Psyche 59: 14-16 (female).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 166. — Scherba, 1961. N. Y. Ent. Soc., Jour. 69:
71-87 (reproduction, nest structure). —Scherba, 1963. N. Y. Ent. Soc., Jour. 71: 219-231.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 599-601. —Gregg, 1963. Ants of Colo.,
pp. 599-601. —Scherba, 1964. N. Y. Ent. Soc., Jour. 72: 231.

Formica ulkei Emery
N. S. w. to Man. s. to Ohio, Ind., 111., Iowa, N. Dak., Wyb. Ecology: The large

mounds may be found in woods or grasslands and occasionally have a thin layer of plant

debris on their surfaces. Host: Formica fusca L.
Formica ulkei Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 653. ♂.
Formica ulkei var. hebescens Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53:

396,487. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 396, 485^487 (each
caste). —Buren, 1944. Iowa State Col, Jour. Sci 18: 300. —Gregg, 1944. Ent. Soc. Amer.,
Ann. 37: 472. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 168 (larva).

Biology: Holmquist, 1928. Ecology 9: 70-87. — Holmquist, 1928. Physiol Zool. 1: 325-327.
—Park, 1929. Psyche 36: 195-215. — Dreyer and Park, 1932. Psyche 39: 127-133.
—Creighton, 1934. Psyche 41: 185-200. —Park, 1936. Psyche 42: 216-231. —Scherba, 1958.
Insectes Sociaux 5: 201-213 (reproduction, mounds). — Kannowski, 1959. Insectes Sociaux
6: 142-143. —Talbot, 1959. Amer. Midland Nat. 61: 128-132 (flight activities). —Scherba,
1959. Amer. Midland Nat. 61: 499-508 (moisture regulation in mounds). —Talbot, 1961.




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Ecology 42: 202-205 (mounds). — Scherba, 1962. Amer. Midland Nat. 67: 373-385 (mound
temperatures). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 256-258.

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).

Species Group Rufa

Nests of members of this group are varied, some species build mounds and others make con-
siderable use of thatching. Nests are usually started beneath objects. Nest foimding females of
some species may behave as temporary social parasites.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 1-10 (N. Amer. variants of Formica
rufa). —Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bui. 104: 472-496.

Formica aterrima Cresson
Colo. Known only from the type male which has not been associated with
any other described form.
Formica aterrima Cresson, 1865. Ent. Soc. Phila., Proc. 4: 426. ♂.

Taxonomy: Brown, 1947. Ent. News 58: 8-9. —Gregg, 1963. Ants of Colo., pp. 544, 546.

Formica calviceps Cole
N. Mex. (Capulin Mt. Natl. Mon.). Ecology: Colony found beneath large stones
which were banked with detritus.
Formica (Formica) calviceps Cole, 1954. Tenn. Acad. Sci., Jour. 29: 164-165. ♂.

Formica ciliata Mayr
Minn., N. Dak., Mont., Wyo., Colo., Utah. Ecology: Nests in meadows or open
woods; sometimes utilizes thatching.
Formica ciliata Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36: 428. ♂.

Taxonomy: Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19: 639-643 (each caste). —Wheeler,
1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392-393, 452-454 (each caste).

Biology: Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 88. —Wheeler, 1910. Ants, pp. 114, 120, 205,
351, 444^145, 450. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 233-234. —Gregg,
1963. Ants of Colo., pp. 545-547.

Formica comata Wheeler
S. Dak., Mont., Colo. Ecology: Nests under stones which are banked with
thatch.
Formica cmnata Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 85. 5, 9, 6.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392, 393, 454^156 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376, 379 (also biological notes).

Biology: Gregg, 1963. Ants of Colo., pp. 547-548, 550.

Formica criniventris Wheeler
N. Dak., S. Dak., Mont. s. to Colo., Utah. Ecology: Found in meadows
and open forests where it nests tmder stones banked with thatch.
Formica crinita Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 87. ♂, ♀. Preocc. by Smith, 1858.
Formica criniventris Wheeler, 1912. Psyche 19: 90. N. name for crinita Wheeler.

Taxonomy: Wheeler 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392, 393, 457-458.
—Cole, 1942. Amer. Midland Nat. 28: 376, 379.

Biology: Wheeler, 1910. Ants, pp. 114, 444-145, 450. —Wheeler and Wheeler, 1944. N. Dak

Hist. Quart. 11: 260. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 234-236. —Gregg,

1963. Ants of Colo., pp. 549-551.

Formica dakotensis Emery
N. S., Ont. w. to Alaska, B. C. s. to Ohio, Iowa, N. Mex., Nev. Ecology:
Usually found in grasslands where it nests in earthen mounds or under stones, about
roots of plants, and in grass clumps banked with considerable detritus. Host: Formica

Formica fusca
***authority mismatch
L., F. lepida Wheeler, F. mxmiana Emery, F. pallidefulva Latreille. Some authors
have recognized two subspecies, dakotensis east of the Rockies and montigemi in the
rocldes and westward.

Formica dakotensis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 644, 652. ♂.

Formica fusca subpolita var. ? specularis Emery, 1893. Zool. Jahrb., Abt. f System. 7:
663. ♀.

Formica dacotensis var. Wasmanni Forel, 1904. Soc. Ent. de Belg., Ann. 48: 153. 5, 9, 6.

Formica montigena Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 374. 5,9, 6.

Formica dakotensis var. saturata Wheeler, 1917. Amer. Acad. Arts and ScL, Proc. 52: 542.
5.




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Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53: 391, 393, 394, 462-465.
— Buren, 1944. Iowa State Col, Jour. Sci. 18: 299. —Brown, 1957. Ent. News 68: 167.

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 364-365. —Wheeler, 1910. Ants, pp.
113, 205, 444-445. —Abbott, 1926. Ent. News 37: 210-211. —Wheeler and Wheeler, 1963.
Ants of N. Dak., pp. 236-238. —Gregg, 1963. Ants of Colo., pp. 551-554.

Morphology: Abbott, 1937. Ent. Soc. Amer., Ann. 20: 117-122 (physiology).

Formica ferocula Wheeler
111. (Rockford). Ecology: Type specimens were found nesting in a dry open
field in small crater nests.
Formica ferocula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53: 392, 461. ♂.

Formica fossaceps Buren
Que., Iowa, N. Dak. Ecology: Found in grasslands and wooded pastures
nesting under stones or fallen logs banked with thatch or in low earthen mounds
covered with fine thatch of grass materials.
Formica fossaceps Buren, 1942. Iowa State Col., Jour. Sci. 16: 402. 5 , 9 , 6.

Biology: Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 302. —King, 1949. Iowa Acad. Sci.,
Proc. 56: 367-370 (mixed colony with F. obscuriventris clivia). — King, 1951. Iowa Acad.
Sci., Proc. 58: 487-489. —King, 1952. Iowa Acad. Sci., Proc. 59: 469-474 (macropseudogynes,
or ? pterergates). — King and Sallee, 1957. Iowa Acad. Sci., Jour. 64: 667-669 (mixed colony
with F. obscuriventris clivia). — King and Sallee, 1959. Iowa Acad. Sci., Proc. 66: 472-473
(slave of F. rubicunda). —King and Sallee, 1962. Iowa Acad. Sci., Proc. 69: 531-539 (mixed
colony with F. obscuriventris clivia). — Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
238-239. — Finnegan, 1973. Canad. Ent. 105: 441-444 (diurnal foraging activity).

Formica haemorrhoidalis Emery
N. Dak., S. Dak., Colo. w. to B. C, Wash., Calif. Ecology: Found in

areas of moderate to sparse cover where nests are usually started under logs or stones

and later with moderate use of thatching.
Formica rufa Integra var. haemorrhoidalis Emery, 1893. Zool. Jahrb., Abt. f. System. 7:

644,652. ♀.
Formica truncicola integroides var. ravida Wheeler, 1913. Harvard Univ., Mus. Comp.

Zool., Bui. 53: 560. ♂, ♀.
Formica truncicola integroides var. tahoensis Wheeler, 1917. Amer. Acad. Arts and Sci.,

Proc. 52: 538. ♂, ♀.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392, 394, 441^42 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376. —Brown, 1965. Ent. News 76: 181-186.
—Cole, 1956. Tenn. Acad. Sci., Jour. 3: 260. —Wheeler and Wheeler, 1963. Ants, of N. Dak,
p. 230 (forms B, C, and D).

Biology: Wheeler, 1910. Ants, pp. 206, 444. — Creighton, 1940. Amer. Mus. Novitates 1055:
6-7. — Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 88-89. —Gregg, 1963. Ants of Colo., pp.
554-557, 579-581.
Integra Nylander. N. S. s. to Ga. w. to Mich., S. Dak., 111., Miss. Ecology: Large colonies are
found in open forests and woods where they nest in stumps and logs or under stones;
plant debris commonly covers the nest.
Formica integra Nylander, 1856. Ann. des Sci. Nat., Zool. 5: 62. ♂.
Formica integra var. similis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 425. 5 , 9 , i.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392, 394, 444^145 (each
caste).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 399. —Wheeler, 1910. Ants, pp.
204-206, 222, 351, 444. —Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bui. 17: 18-19.
—Creighton, 1940. Amer. Mus. Novitates 1055: 6. — Kloft, et al., 1973. Fla. Ent. 56: 67-76
(habitat, nest structure, polygamy, biometry).

Formica integroides
***authority mismatch
coloradensis Wheeler. Colo., N. Mex., Idaho, Utah. Ecology: Found in forests at
high elevations (mostly 8,000 to 10,000 feet); nests are started under stumps or logs and
later may be large and dome-shaped with extensive use of thatch.
Formica truncicola integroides var. coloradensis Wheeler, 1913. Harvard Univ., Mus.
Comp. Zool., Bui. 53: 393, 440. ♂, ♀.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 377, 381.




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Biology: Creighton, 1940. Amer. Mus. Novitates 1055: 6-7. —Gregg, 1963. Ants of Colo. pp.
557-560.
integ^oides integroides Emery. Wash., Oreg., Calif. Ecology: Inhabits open woods in the
coastal mountains and west slopes of the Cascades and Sierras where it nests under
logs and stumps banked with plant debris.
Formica rufa obscuriventris var. integroides Emery, 1893. Zool. Jahrb., Abt. f. System. 7:

644, 649. ♂.
Formica truncicola integra var. subcaviceps Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:540. 5, cJ.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 393, 394, 438-439.
—Creighton, 1940. Amer. Mus. Novitates 1055: 6-7.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 537. —Miller, 1957. Insectes
Sociaux 4: 253.

Formica integroides
***authority mismatch
planipilis Creighton. N. Dak., Colo., Nev. Ecology: Found in grasslands and open
forests; nests are frequently started at bases of plants and extensive use is made of
thatching; completed nests are moundlike.
Formica rufa planipilis Creighton, 1940. Amer. Mus. Novitates 1055: 7, 9. ♂.

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31: 259 (female).

Biology: Wheeler and Wheeler, 1963. Ants, of N. Dak., pp. 248-249. —Gregg, 1963. Ants of
Colo., pp. 560-561. —Cole, 1966. Brigham Young Univ., Sci. Bui., Biol. Ser. 7: 23 (Nev. Test
Site).

Formica integroides
***authority mismatch
propinqua Wheeler. Colo., Utah, Wash., Calif. Ecology: Found on the east slopes
of the Cascades and Sierras in Wash, and Calif.; inhabits forests where nests are
started under logs and stumps and considerable use is made of thatching.
Formica truncicola integroides var. propinqiui Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:538. ♀.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 6-7. —Cole, 1956. Tenn. Acad. Sci.,
Jour. 31: 267-259 (female, male).

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 88. —Gregg, 1963. Ants of Colo., pp.
562-564.

Formica integroides
***authority mismatch
subfasciata Wheeler. Calif. (Mill Creek Canyon, Wilson Peak, 7500 ft., San
Bernardino Mtns.).
Formica truncicola integroides var. subfasciata Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:539. ♀.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 7. — Brovra, 1965. Ent. News 76:
185 (possible syn. of integroides Emery).

Formica laeviceps Creighton
N. Dak., S. Dak., Colo., Utah. Ecology: Nests mostly in open areas under
stones and logs which are banked vnth little debris.
Formica rufa laeviceps Creighton, 1940. Amer. Mus. Novitates 1055: 7, 9. ♂, ♀.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 377, 380 (also biological notes).

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 239-240. —Gregg, 1963. Ants of
Colo., pp. 564-565.

Formica lugubris Zetterstedt
Que.; Europe. Ecology: Populous colonies are found in large mound nests
of thatch and debris in open forests. Introduced into Valcartier, Quebec in 1971 from
Italy for use as a predator of the Swaine jack pine sawfly, Neodiprion swainei Midd. It
is apparently established in Quebec.
Formica lugubris Zetterstedt, 1840. Ins. Lapponica, v. 1, p. 449. ♂.

Taxonomy: Yarrow, 1955. Soc. Brit. Ent., Trans. 12: 5, 10 (synonymy, also biological notes).

Biology: Pavan, 1959. Min. deU' Agr. e delle For. [Italy] 4: 5 (in Italy). — Pavan, 1963. Pavia
Univ. Symp. Genet, et Biol. Ital. 11: 61-84 (use to protect alpine forests against insect
damage). —Pavan, 1963. Pavia Univ. Symp. Genet, et Biol. Ital. 12: 122-131 (reconstruction
of nest). — Klimetzek, 1970. Ztschr. f. Angew. Ent. 66: 84-95 (environmental factors and




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distribution). —Williamson, 1973. Canad. Dept. Agr., Liberation Bui. no. 36, p. 2. (in Que.).
— Finnegan, 1975. Canad. Ent. 107: 1271-1274 (introduction into eastern Canada).

Formica mucescens Wheeler
Colo., Utah. Ecology: Found in open areas where it nests under stones;

moderate use is made of thatching.
Formica truncicola mucescens Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53:

392,394,442. 5, 9,
Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 6-7. — Cole, 1942. Amer. Midland
Nat. 28: 376, 380.

Biology: Gregg, 1963. Ants of Colo., pp. 566-567.

Formica obscuripes Forel
Que.; Mich., Ind., Man. w. to B. C. s. to N. Mex., Utah., Calif. Ecology: Nests
are found in open areas where they are usually started at the base of a plant. Extensive
use is made of thatching and the finished nest is a large mound of detritus. Probably
the most common thatching ant of the western states. Introduced into Quebec from
Manitoba for use as a predator of the Swaine jack pine sawfly, Neodiprkm stuainei
Midd.

Formica rufa obscuripes Forel, 1886. Soc. Ent. de Belg., Ann. (C. R.) 30: 39. ♂.

Formica rufa obscuriventris var. melanotica Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
644,650. ♂.

Formica rufa obscuriventris var. rubiginosa Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
644, 650. "♀"=♂. Preocc. by Guillou, 1841.

Formica rufa a^gerans Wheeler, 1912. Psyche 19: 90. N. name for rubiginosa Emery.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53: 392, 394, 432-433 (each
caste). — Creighton, 1940. Amer. Mus. Novitates 1055: 7. — Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 300. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 165-166 (larva).

Biology: Treheme, 1915. Canad. Ent. 47: 104. — Burrill and Smith, 1919. Ohio Jour. Sci. 19:
286. —Cole, 1932. Psyche 39: 30-33. —Weber, 1935. Ecol. Monog. 5: 165-206 (in N. Dak.).
— Essig, 1926. Ins. West. No. Amer., p. 867. — Eckert and Mallis, 1937. Calif. Agr. Expt.
Sta. Cir. 342: 30. —Weber, 1941. Canad. Ent. 73: 140-141. —Cole, 1942. Amer. Midland Nat.
28: 380. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 263-266. — SaUee and King,
1947. Iowa Acad. Sci., Proc. 54: 349-352. —King and Walters, 1950. Iowa Acad. Sci., Proc.
57: 469-47. —King and Sallee, 1953. Iowa Acad. Sci., Proc. 60: 656-659. —King and Sallee,
1957. Iowa Acad. Sci., Proc. 63: 721-723. —Talbot, 1959. Amer. Midland Nat. 61: 124-128
(flight activities). —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 240-244. —Gregg,
1963. Ants of Colo., pp. 566-572. —Bradley, 1972. Canad. Ent. 104: 245-249 (transplanting
colonies in jack pine stands in Man.). — Talbot, 1972. Kans. Ent. Soc., Jour. 45: 254-258
(flights and swarms). — Alpert and Akre, 1973. Ent. Soc. Amer., Ann. 66: 753-760
(association with inquiline, Leptothorax diversipilosvs Smith, in Wash.). — Williamson,
1973. Canad. Dept. Agr., Liberation Bui. no. 36, p. 2 (introduction into Que.).

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).

Formica obscuriventris
***authority mismatch
clivia Creighton. Man., Wis., Iowa w. to B. C. s. to N. Mex., Utah. Ecology:
Nests in forests under logs and stones; moderate use is made of thatching.
Formica rufa clivia Creighton, 1940. Amer. Mus. Novitates 1055: 8, 9. 5, 9, S.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 377. —Buren, 1944. Iowa State Col., Jour. Sci.
18: 300, 302.

Biology: King, 1949. Iowa Acad. Sci., Proc. 56: 367-370 (mixed colony with F. fossaceps
Buren). —King and Sallee, 1951. Iowa Acad. Sci., Proc. 58: 487^189. —King, 1955. Iowa
Acad. Sci., Proc. 62: 509-513 (winged workers). —King and SaUee, 1959. Iowa Acad. Sci.,
Proc. 66: 472^73 (slave of F. rubicunda Emery). —King and Sallee, 1962. Iowa Acad. ScL,
Proc. 69: 531-539 (mixed colonies with F. fossaceps Buren). —Gregg, 1963. Ants of Colo.,
pp. 574-577.

Formica obscuriventris
***authority mismatch
obscuriventris Mayr. Que., Maine s. to Va. w. to N. Dak., Iowa, Colo. Ecology:
Nests in woods and grasslands under logs and makes moderate use of thatching.
Formica truncicola var. obscuriventris Mayr, 1870. Zool.-Bot. (jJesell. Wien, Verb. 20: 951.

5.
Formica dryas Wheeler, 1905. Amer. Mus. Nat. Hist, Bui. 21: 268. P, 9.




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Formica dryas var. gymnomma Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 269. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392, 448. — Creighton,
1940. Amer. Mus. Novitates 1055: 8.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24: 623. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bui. 22: 595, 597. — Buren, 1941. Iowa State Col, Jour. Sci. 15:
115. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 244-246. —Gregg, 1963. Ants of
Colo., pp. 572-574. —Talbot, 1964. Anim. Behaviour 12: 154 (nest structure and
flights).— Ayre, 1968. Canad. Ent. 100: 165-172 (prey finding, capture and transport).

Formica oreas
***authority mismatch
comptula Wheeler. Sask., N. Dak., Iowa w. to Alta., Wash., Idaho, Utah. Ecology: Nests
in wooded areas or grasslands under stones or logs banked with detritus.
Formica oreas var. comptula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53:
391, 393, 460. ? , 9 .

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 376, 380.

Biology: Cole, 1934. Psyche 41: 227. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp.
246-248.

Formica oreas
***authority mismatch
oreas Wheeler. Mont., Wyo., Colo., N. Mex., Utah. Ecology: Nests in open woods or

meadows under rocks or logs covered with detritus. Colonies may be abundant and have
sizeable mounds.
Formica oreas Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19: 643. 5, 9,6.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 391, 393, 458^160.
—Cole, 1942. Amer. Midland Nat. 28: 376, 379.

Biology: Wheeler, 1910. Ants, pp. 114, 205, 351, 444. —Gregg, 1963. Ants of Colo., pp. 577-579.

Formica prociliata Kennedy and Dennis
Ohio, Wis., Iowa. Ecology: Nests of the type specimens were
under stones on ledges in open woods and lacked thatching.
Formica prociliata Kennedy and Dennis, 1937. Ent. Soc. Amer., Aim. 30: 531. 5, 9, 6.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 303.

Formica reflexa Buren
Que., Minn., Iowa, N. Dak Ecology: Has only been found in association with the
host; a permanent Lnquiline ?. Host: Formica fusca L.
Formica reflexa Buren, 1942. Iowa State Col., Jour. Sci. 16: 399. 5, 9, 6.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 299, 303.

Biology: King, 1949. Iowa Acad. Sci., Proc. 56: 367-379 (mixed colony vdth F. fusca). —King,
1951. Iowa Acad. Sci., Proc. 58: 487^489. —Wheeler and Wheeler, 1963. Ants of N. Dak, p.
249-250. — Francoeur and Beique, 1968. Nat. Canad. 95: 228 (Que.).

Formica subnitens Creighton
N. Dak, Wyo., Colo., N. Mex., B. C, Oreg., Calif. Ecology: Usually nests
in grasslands, under stones banked with debris or in dome-shaped moimds of thatch or
detritus.
Formica rufa subnitens Creighton, 1940. Amer. Mus. Novitates 1055: 7, 10. ♂.

Taxonomy: Cole, 1955. Tenn. Acad. Sci., Jour. 30: 50. —Miller, 1957. Insectes Sociaux 4:
253-257 (each caste).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 165. —Ayre, 1957. Insectes Sociaux 4: 173-176
(ecological notes). -Ayre, 1958. Insectes Sociaux 5: 1-7. —Ayre, 1958. Insectes Sociaux 5:
147-157 (meterological factors affecting foraging). —Ayre, 1959. Insectes Sociaux 6:
105-114 (food habits). —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 250-252. —Gregg.
1963. Ants of Colo., pp. 581-584.

Species Group Microgyna

Species of this group are believed to be temporary social parasites of other species of For-
mica. The female in some way is adopted by workers of the host species. Host workers may
remain in the colony after the intruding queen has established her own brood, but the host wor-
kers eventually die. Most species are found in open woods or meadows. The nests are usually of
the thatch type, but the thatching is normally scattered about the nest openings and appears as
a flattened disc.




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Taxonomy: Letendre and Huot, 1972. Soc. Ent. Quebec, Ann. 17: 117-132 (preliminary
considerations to a revision of the microgyna group).

Formica densiventris Viereck
Colo., N. Mex., Utah w. to CaUf. Ecology: Found in forests and open
forests, under stones and logs; only occasionally with use of thatch.

Formica fusca var. densiventris Viereck, 1903. Amer. Ent. Soc, Trans. 29: 74. ♂.

Formica microgyna rasilis var. spicata Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bui. 53: 395, 469. 5, 9, d.

Formica microgyna rasilis var. pinetorum. Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 542. ♂.

Taxonomy: Brown, 1947. Ent. News 58: 6-8. —Cole, 1955. Tenn. Acad. Sci., Jour. 30: 50.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bui. 40: 85. —Cole, 1954. Tenn. Acad. Sci., Jour.
29: 165. —Gregg, 1963. Ants of Colo., pp. 589-593.

Formica difficilis Emery
Mass. s. to Ga. w. to Iowa. Ecology: Nests are in woodlands, sometimes under
stones, and are packed with vegetable debris. Host: Formica pallidefulva (Latreille).
Formica rufa difficilis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 651. 3, 9, 6.
Formica difficilis var. consocians Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 371. ♀.

2, Formica habrogyna Cole, 1939. Amer. Midland Nat. 22: 413. ♂, ♀.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 395, 477-480 (each
caste). — Buren, 1944. Iowa State Col, Jour. Sci. 18: 306. —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 46: 164 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 347-373. —Wheeler, 1906. Amer. Mus.
Nat. Hist., Bui. 22: 50-64. —Wheeler, 1910. Ants, pp. 113-114, 205-206, 441-444.

Formica dirksi Wing
Maine (Daigle, Aroostook Co.). Ecology: The host nest of the type queen was at
the edge of a clearing under bark of a dry stump. Host: Formica fusca L.
F(yrmica dirksi Wing, 1949. Canad. Ent. 81: 13-15. ♀.

Formica impexa Wheeler
Mass., Mich., Minn. Host: Formica fusca L.

Formica impexa Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 273. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp Zool., bul. 53: 395-396, 472-473
(worker, female).

Biology: Wheeler, 1906. Psyche 13: 40. —Wheeler, 1910. Ants, pp. 113, 444.

Formica indianensis Cole
Ind., Iowa. Ecology: Nest was in a grassy field under a stone and was
surrounded with grass debris.
Formica indianensis Cole, 1940. Amer. Midland Nat. 23: 224. 5 , (5 .

Taxonomy: Buren, 1944. Iowa State Col, Jour. Sci. 18: 300, 306.

Formica knighti Buren
Iowa (Bonaparte). Ecology: Nest was in a pasture under bushes and
surmoimted by a low dome of plant debris.
Formica (Formica) knighti Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 303-305. ♂.

Formica microgyna Wheeler
Wyo., Colo., N. Mex., Utah, Nev. Ecology: Prefers meadows and open
forests. Nests are usually started under stones which are piled with thatching, the
thatching increasing in size as the nest g^ws. Host: Formica argentea Wheeler, F.

Formica lasioides Emery
F. neogagates Emery.
Formica microgyna Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19: 645. 5, 9, 6.
Formica m,icrogyna microgyna var. reddiva Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 395, 467. 5,
Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., BuL 53: 395-396, 465-467 (each
caste).

Biology: Wheeler, 1917. Amer. Acad. Arts and ScL, Proc. 52: 542. —Gregg, 1963. Ants of
Colo., pp. 584-587. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 24 (Nev. Test
Site).
morsel Wheeler. Mass. (South Natick). ^

Formica m/yrsei Wheeler, 1906. Psyche 13: 39. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp.ZooL, Bul. 53: 394, 480-481.




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Formica nepticula Wheeler
Mass., Conn., 111., Iowa. Ecology: Nests in open woods under stones or
rotting limbs and are banked with vegetable debris.
Formica nepticula Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 270. 9, 9, S.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 394, 396, AlbAll (each
caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22: 595, 597.
— Buren, 1944. Iowa State Col., Jour. Sci. 18: 30, 306.

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 64. —Wheeler, 1910. Ants, pp. 113,
205, 444.

Formica nevadensis Wheeler
Nev. (Ormsby Co.). Ecology: Nests in open forests under stones and
surmounted by plant debris.
Formica microgyria var. nevadensis Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20: 373. ♀.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bui. 21: 272. —Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool., Bui. 53: 396, 470-472 (worker). —Wheeler, 1917. Amer. Acad. Arts
and Sci., Proc. 52: 544. —Cole, 1956. Tenn. Acad. Sci., Jour. 31: 256-257 (also biology).

Biology: Wheeler, 1910. Ants, pp. 11, 444.

Formica postoculata Kennedy and Dennis
Pa., Ind., 111., Iowa. Ecology: Nest of type series from
Indiana was in a pasture between two boulders and was covered with trash.
Formica postoculata Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 540-542. ♂.

Formica querquetulana Kennedy and Dennis
New England to Ohio. Ecology: Nests found in pastures
and sandy areas usually beneath objects and commonly covered with loose trash. Host:
Formica fusca L.
Formica querqiuetulana Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 536-540. ♂.
9.

Formica rasilis Wheeler
Colo., N. Mex., Utah northwest to Wash. Ecology: Nests in open or semi-open
areas usually under stones. Host: Formica argentea Wheeler, F. fusca L. Creighton
(1950) stated that there is a great deal of intergradation between rasilis and spicata
(see synonymy of densiventris); consequently, rasilis and densiventris may be
conspecific.
Formica microgyna var. rasilis Wheeler, 1903. Amer. Mus. Nat. Hist., Bui. 19: 648. 2,9,
cJ.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 395-396, 46&469.
—Creighton, 1950. Harvard Univ., Mus. Comp. Zool, Bui. 104: 506-507.

Biology: Cole, 1942. Amer. Midland Nat. 28: 376, 381. —Gregg, 1963. Ants of Colo., pp.
587-589.

Formica scitula Wheeler
Ga. (Clayton).

Formica microgyna scitula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 395,
470. ♀.

Formica spatulata Buren
Minn., Iowa, N. Dak., Mont., B. C. Ecology: Nests under stones. Host:
Formica fusca L.
Formica (Formica) m.icrogyna spatulata Buren, 1944. Iowa State Col., Jour. Sci. 18: 300,
305. 5, 9, 6.

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., p. 253.

Formica whymperi
***authority mismatch
adamsi Wheeler. Mich, to Minn. Ecology: Has been found in drier tamarack
swamps in nests covered with leaves.
Formica adamsi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 84. ♂.
Formica adam^ Wheeler, 1909. Mich. Geol. Survey, Bui. 5: 326. ♀.

Taxonomy: Wheeler, 1913. Harvard Univ. Mus. Comp. ZooL, BuL 53: 395, 473^74. —Wheeler,

1917. Amer. Acad. Arts and Sci., Proc. 52: 544.
Biology: Drooz, 1960. U. S. Dept. Agr., Tech. Bui. 1212, p. 36 (predator of larch sawfly).

Formica whymperi
***authority mismatch
alpina Wheeler. Colo., N. Mex., Idaho, Utah. Ecology: Found at high elevations,

mostly above 9,000 feet where it nests in forests under stones or rotting logs and with
some plant debris or thatch around these objects. Host: Formica neorufibarbis Emery.
Formica adamsi var. alpina Wheeler, 1909. N. Y. Ent. Soc, Jour. 17: 85. ♂.




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Formica adamsi var. alpina Wheeler, 1909. Mich. Geol. Survey, Bui. 5: 327. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 395, 475. —Wheeler,
1917. Amer. Acad. Arts and Sci., Proc. 52: 544.

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 165-166. —Gregg, 1963. Ants of Colo., pp.
593-596 (also description of female).

Formica whymperi
***authority mismatch
califomica Wheeler. Mont., Idaho., Wash. s. to Calif.

Formica microgyria califomica Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 543.

Formica microgyria califomica var. hybrida Wheeler, 1917. Amer. Acad. Arts and Sci.,

Proc. 52:543. ♂.

Formica whymperi
***authority mismatch
whymperi Forel. Mont. w. to B. C, Wash. Ecology: Nests under stones and logs

which it banks with plant debris. Host: Formica neoclara Emery.
FoTnnica rufa obscuripes var. whymperi Forel, 1904. Soc. Ent. de Belg., Ann. 48: 152. ♂.
Formica microgyria rasilis var. pullula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,

Bui. 53:562. ♂, ♀.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 392, 434. —Wheeler,
1917. Amer. Acad. Arts and Sci., Proc. 52: 544 (also biological notes).

Species Group Sanguinea

Members of this subgenus are social parasites. Nest founding females forceably enter small
colonies of the host species and kill or drive off the host workers before rearing the host brood
for her own use. Hosts are other species of Formica. Some species are known to regularly con-
duct slave raids, a habit unknown in other Formica species.

Taxonomy: WUson and Brown, 1955. Psyche 62: 108-129. — Buren, 1968. Ga. Ent. Soc., Jour.
3: 25-40 (key to spp.).

Biology: Wheeler, 1910. Ants, pp. 458^70. — Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bui. 104: 460-461.

Formica creightoni Buren
Mich., IlL, Iowa. Host: Formica neogagates Emery.

Formica (Raptiformica) creightoni Buren, 1968. Ga. Ent. Soc., Jour. 3: 35-36. ♀.

Formica curiosa Creighton
B. C, Mont., Idaho, Oreg. Host: Formica lasioides Emery, F. manni
Wheeler.
Formica curiosa Creighton, 1935. Amer. Mus. Novitates 773: 5. 2 , 9 .
Formica parcipappa Cole, 1946. Ent. Soc. Amer., Ann. 36: 616. ♂.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 123-125. —Buren, 1968. Ga. Ent. Soc., Jour.
3: 32-33. — Snelling, 1969. Ent. Soc. Wash., Proc. 71: 194-196 (syn. of parcipappa; biological
notes).

Formica emeryi Wheeler
Colo. (Bradmoor, Colorado Springs). Host: Formica neogagates Emery.

Formica emeryi Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 389, 419. ♂, ♀.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 118 (a syn. of subnuda). — Gregg, 1963.
Ants of Colo., pp. 606-608 (valid sp.). —Buren, 1968. Ga. Ent. Soc., Jour. 3: 34-35 (valid sp.).

Formica pergandei Emery
Que., N. H. s. to N. C. w. to N. Dak., S. Dak., Iowa, Colo. Host: Formica

Formica fusca
***authority mismatch
L., F. pallidefulva Latreille (?).
Formica pergandei Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 646. ♀.
Formica sanguinea rubicunda var. subluxnda Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bui. 53: 390, 408. 9, 9.

Taxonomy: Buren, 1968. Ga. Ent. Soc., Jour. 3: 28-29.

Biology: Wheeler, 1901. Amer. Nat. 35: 722. —Gregg, 1946. Amer. Midland Nat. 35: 752.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 219-220. —Gregg, 1963. Ants of Colo.,
pp. 622-623. — Regnier and Wilson, 1971. Science 172: 267-269 (chemical communication).
—Hung, 1973. Ent. News 84: 253-259 (reproductive biology). — Finnegan, 1973. Canad.
Ent. 105: 441-444 (diurnal foraging activity; as sublucida).




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Formica puberula Emery
Man., N. Dak., S. Dak., Wyo., Colo., Tex. w. to B. C, Wash., Calif. Host:

Formica altipetens Wheeler, F. bradleyi Wheeler, F. fusca L., F. hevntti Wheeler, F.

Formica lasioides Emery
F. lepida Wheeler, F. montana Emery, F. neoclara Emery, F.

Formica neorufibarbis Emery
F. pallidefulva nitidiventris Emery, F. rasilis Wheeler, F.

Formica subpolita Mayr

Formica sanguinea puberula Emery, 1893. Zool. Jahrb., Abt. f. System 7: 643, 648. ♀.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 390, 413^14 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376. — Buren, 1968. Ga Ent. Soc., Jour. 3:
30-31.

Biology: Wheeler, 1910. Ants, pp. 458-460. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
220-221. —Gregg, 1963. Ants of Colo., pp. 613-616. — SnelUng, 1969. Ent. Soc. Wash., Proc.
71: 196-197.

Formica rubicunda Emery
Ont. s. to N. C, Tenn. w. to Mont., Colo., N. Mex. Host: Formica altipetens
Wheeler, F. bradleyi Wheeler, F. fossaceps Buren, F. fusca L., F. lasioides Emery, F.

Formica lepida Wheeler
F. montana Emery, F. neoclara Emery, F. neogagates Emery, F.

Formica neorufibarbis Emery
F. obscuriventris clivia Creighton, F. pallidefulva nitidiventris
Emery, F. schaufussi Mayr.
Formica sanguinea rubicunda Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 647. 9,9.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool, Bui. 53: 390, 406^08 (each
caste). —Wheeler, 1917. Conn. State (Jeol. and Nat. Hist. Survey Bui. 22: 595. —Buren,
1944. Iowa State Col., Jour. Sci. 18: 299. —Buren, 1968. Ga. Ent. Soc., Jour. 3: 30.

Biology: Wheeler, 1910. Ants, pp. 45^470. —Smith, 1928. N. Y. Ent. Soc., Jour. 26: 327-329.
—Talbot, 1934. Ecology 15: 421, 430-434. —King and SaUee, 1959. Iowa Acad. Sci., Proc. 66:
472-473. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 221-223. —Gregg, 1963. Ants
of Colo., pp. 616-618.
subintegrra Emery. Newfoundland, N. S., Ont. s. to S. C, Tenn. w. to N. Dak, Iowa, Kans.

Host: Formica fusca L., F. lasioides Emery, F. montana Emery, F. neogagates Emery,

F. pallidefulva nitidiventris Emery, F. schaufussi Mayr, F. subpolita Mayr.
Formica sanguinea rubicunda var. subintegra Emery, 1893. Zool. Jahrb., Abt. f. System.

7: 643, 648. ♂.
Formica sanguinea subintegra var. gilvescens Wheeler, 1913. Harvard Univ., Mus. Comp.

Zool., Bui. 53: 390, 412. ♂.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 390, 410-412.
—Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bui. 22: 595. —Buren, 1944.
Iowa State Col., Jour. Sci. 18: 299. —Wilson and Brown, 1955. Psyche 62: 120. —Buren,
1968. Ga. Ent. Soc., Jour. 3: 29-30.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist, Bui. 24: 624, 627-631. —Smith, 1928. N. Y.
Ent. Soc. Jour. 36: 323-327. —Talbot and Kennedy, 1940. Ent. Soc. Amer., Ann. 33: 560-577.
—Brown, 1958. Psyche 65: 39-40. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
225-227. — Kannowski, 1963. Pavia Univ., Symp. Genet. Biol. Ital. 12: 74-102 (mating
behavior). — Regnier and Wilson, 1971. Science 172: 267-269 (chemical communication).

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).

Formica subnuda Emery
Newfoundland w. to Yukon, Alaska s. to N. Y., Minn., N. Dak., Colo., N. Mex.,
Ariz., Calif. Host: Fcrrmica altipetens Wheeler, F. fusca L., F. numtana Emery, F.

Formica neorufibarbis Emery
F. subpolita Mayr.
Formica sanguinea rubicunda var. subnuda Emery, 1895. Zool. Jahrb., Abt. f. System. 8:

335. ♂.
Formica sanguinea aserva Forel, 1901. Soc. Ent. de Belg., Ann. 45: 395. ♂, ♀.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bui. 53: 389, 404-406, 409-410
(each caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist Survey Bui. 22: 595.
—Buren, 1944. Iowa State CoL, Jour. Sci. 18: 299, 308. —Wheeler and Wheeler, 1953. Ent.
Soc. Amer., Ann. 46: 170 flarva). —Wilson and Brovm, 1955. Psyche 62: 118. —Buren, 1968.
Ga. Ent. Soc., Jour. 3: 33-34.




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Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bui. 22: 85. —Wheeler, 1908. Amer. Mus. Nat.
Hist, Bui. 24: 623, 631-633. —Wheeler, 1910. Ants, pp. 458-460, 468. —Wheeler, 1917.
Harvard Univ., Mus. Comp. Zool., Bui. 61: 19. —Cole, 1942. Amer. Midland Nat. 28: 378.
—Gregg, 1947. Colo. Univ. Studies 2: 393. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 223-225. —Gregg, 1963. Ants of Colo., pp. 618-621. —Chapman, 1957. Canad. Ent. 89:
392 (swarming on mountain tops). — Finnegan, 1973. Canad. Ent. 105: 441-444 (diurnal
foraging activity).

Formica wheeleri Creighton
N. Dak., Colo., N. Mex., Utah, Ariz. Host: Formica altipetens Wheeler, F.

Formica bradleyi Wheeler
F. fusca L., F. lasioides Emery, F. lepida Wheeler, F. neogagates

Emery, F. neorufibarbis Emery.
Formica wheeleri Creighton, 1935. Amer. Mus. Novitates 773: 1-5. ♂, ♀.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 125-126. — Buren, 1968. Ga. Ent. Soc., Jour.
3: 31-32.

Biology: Cole, 1942. Amer. Midland Nat. 28: 376, 378. —Cole, 1954. Tenn. Acad. Sci., Jour. 29:
104. — WUson, 1955. Psyche 62: 130-133. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 227-228. —Gregg, 1963. Ants of Colo., pp. 623-625. — Halverson, et al., 1976. Kans. Ent.
Soc, Jour. 49: 299 (bradleyi as slave).

Genus POLYERGUS Latreille

Polyergus Latreille, 1804. Nouv. Diet. Hist. Nat., v. 24, p. 179.
Type-species: Formica rufescens Latreille. Monotypic.

The species of this genus are obligatory or true slave-making ants. In nest founding, the
female enters a nest of the host species, eventually kills the rightful queen, and uses the host
workers to tend her brood. A colony of Polyergus will conduct slave raids on nests of species of
Formica, and workers of the host are taken and used by the Polyergtis colony to feed and rear
the brood and excavate the nest.

Revision: Smith, 1947. Amer. Midland Nat. 38: 150-161.

Taxonomy: Wheeler, 1968. Ent. Soc. Wash., Proc. 70: 156-164 (male genitalia and taxonomy).

Morphology: Forbes and Brassel, 1962. N. Y. Ent. Soc., Jour. 70: 79-87 (male genitalia and
terminal segments).

Polyergus breviceps Emery
Ont., Mich. w. to B. C. s. to Ind., 111., Mo., Kans., N. Mex., Ariz., Calif. Host:

Formica altipetens Wheeler, F. argentea Wheeler, F. fusca L., F. lepida Wheeler, F.

montana Emery, F. neoclara Emery, F. neorufibarbis Emery, F. pallidefulva

nitidiventris Emery, F. schaufussi schaufussi Mayr, F. subpolita Mayr.
Polyergtis rufescens breviceps Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 666. ♂.
Polyergus rufescens bicolor Wasmann, 1901. Allg. Ztschr. f. Ent. 6: 369. 5, 9, 6.
Polyergus rufescens breviceps var. Silvestrii Santschi, 1909. Soc. Ent. Ital., Bol. 41: 7. ♂.

6.
Polyergus rufescens breviceps var. montezuma Wheeler, 1914. N. Y. Ent. Soc., Jour. 22: 56.

5, 9,cJ.
Polyergus rufescens breviceps var. umbratus Wheeler, 1915. Amer. Mus. Nat. Hist., Bui.

34:419. ♂.
Polyergus rufescens laeviceps Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 420. ♀.
Polyergus rufescens breviceps var. fusciventris Wheeler, 1917. Amer. Acad. Arts and Sci.,

Proc. 52: 555. ♂.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 385. —Buren, 1944. Iowa State CoL, Jour. Sci.
18: 310. —Smith, 1947. Amer. Midland Nat. 38: 152, 157-159, 161. —Wheeler, 1968. Ent.
Soc. Wash., Proc. 70: 156-164. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 214
(larva, as rufescens).

Biology: Wheeler, 1910. Ants, pp. 475^182. —Wheeler, 1916. N. Y. Ent. Soc., Jour. 24: 107-118.
—Smith, 1928. N. Y. Ent. Soc., Jour. 36: 329-333. — Mallis, 1941. South. Calif. Acad. Sci.,
Bui. 40: 82. —Wheeler and Wheeler, 1944. N. Dak Hist. Quart. 11: 260. —Gregg, 1946.
Amer. Midland Nat. 35: 754. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 275-277.




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—Gregg, 1963. Ants of Colo., pp. 635-643. —Hung, 1973. Ent. News 84: 253-259
(reproductive biology).

Polyergus lucidus
***authority mismatch
longicornis Smith. N. C, S. C, Ga. Host: Formica scha2ifussi schaufussi Mayr.

Polyergus lucidus longicornis Smith, 1947. Amer. Midland Nat. 38: 155. ♀.

Polyergus lucidus
***authority mismatch
lucidus Mayr. Mass. s. to S. C. w. to N. Dak, Iowa, Colo., N. Mex. Host: Formica
lasioides Emery, F. neogagates Emery, F. pallidefulva nitidiventris Emery, F.
schaufussi schaufussi Mayr.
Polyergus lucidus Mayr, 1870. Zool.-Bot. Gesell. Wien, Verb. 20: 952. ♂.
Polyergus luxndus montivagus Wheeler, 1915. Amer. Mus. Nat. Hist., Bui. 34: 419. ♂, ♀.
6.

Taxonomy: Mayr, 1886. Zool.-Bot. Gesell. Wien, Verb. 36: 424. — Buren, 1944. Iowa State
Col, Jour. Sci. 18: 310. —Smith, 1947. Amer. Midland Nat. 38: 152-156. —Wheeler and
Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 214 (larva).

Biology: McCook, 1880. Acad. Nat. Sci. Phila., Proc. 32: 376-384. — Burrill, 1908. N. Y. Ent.
Soc., Jour. 16: 144-151. —Wheeler, 1910. Ants, pp. 482-486. —Smith, 1934. N. Y. Ent. Soc.,
Jour. 42: 360-361. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 274-275. —Gregg,
1963. Ants of Colo., pp. 633-635. —Talbot, 1967. Psyche 74: 299-313 (slave raids). —Talbot,
1968. Psyche 75: 46-52 (flights). — Harman, 1968. Ent. News 79: 217-223 (ecology, near
Chicago). — MarUn, 1968. 111. State Acad. Sci., Trans. 61: 207-209 (colony formation).
— Marlin, 1969. Kans. Ent. Soc, Jour. 42: 108-115 (raiding behavior). — Marlin, 1971. Amer.
Midland Nat. 86: 181-189 (mating, nesting, and ant enemies).

Unplaced Taxa of Formicinae

Formica arenicola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 160. 5 . Washington, D. C.
Formica atra Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. 5 . Washington, D. C.
Formica Connecticutensis Buckley, 1866. Ent. Soc. Phila., Proc. 6: 154. 5,9. Conn., N. Y., D. C.
Formica lauta Say, 1836. Boston Jour. Nat. Hist. 1: 286. ♀, ♂.
Formica monticola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 157. 5, 9, cJ. N. Y.
Formica Nortonii Buckley, 1866. Ent. Soc. Phila., Proc. 6: 153. 5,9. Conn.
Formica Nova Anglae Buckley, 1866. Ent. Soc. Phila., Proc. 6: 153. 5,9. Maine.
Formica occidentalis Buckley, 1866. Ent. Soc. Phila., Proc. 6: 157. 5,9. Conn., N. Y.
Formica politurata Buckley, 1866. Ent. Soc. Phila., Proc. 6: 160. 5 . Mich.
Formica saxicola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 166. 5,9. Buchanan Co., Tex.
Formica sem,ipunctata Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 262. 9 . "Taken on a journey
from New York to Cumberland House."
Formica septentrionale Buckley, 1866. Ent. Soc. PhUa., Proc. 6: 161. 5,9. Mich., 111.
Formica tenuissima Buckley, 1866. Ent. Soc. Phila, Proc. 6: 159. 5. Central Tex.
Formica Virginiana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. 5. Washington, D. C.

Unplaced Taxa of Formicidae

Formica dislocata Say, 1836. Boston Jour. Nat. Hist. 1: 288. ♀, ♂.
Formica Lincecumii Buckley, 1866. Ent. Soc. Phila., Proc. 6: 163. 5, 9, cJ. Tex.
Polyergus Texana Buckley, 1866. Ent. Soc. Phila, Proc. 6: 170. 9. Buchanan Co., Tex.
Formica triangularis Say, 1836. Boston Jour. Nat. Hist. 1: 288. ♀, ♂.



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