Wheeler, George Carlos (1897-1991)

BIOGRAPHICAL NOTE
George Carlos Wheeler (1897-1991) went to Texas Christian University in 1914 with the intention of becoming a high school language teacher. His long-time interest in biology, however, dominated his spare time and upon transferring to the Rice Institute in 1915, Wheeler began to enroll in biology classes. His first teacher and academic advisor was Julian Huxley, who strongly influenced Wheeler to become a professional biologist. When Wheeler completed his undergraduate work at the Rice Institute in 1918, he enlisted in the army and was sent to the Yale Army Laboratory School. After Armistice, he carried out his military service at Base Hospital in Camp Dodge, Iowa. In 1919, he started his graduate work in entomology at Harvard's Bussey Institute. After he received his PhD, Wheeler went on to teach at Syracuse University, the University of North Dakota and the Desert Research Institute (University of Nevada). In 1986, Wheeler authored a brief autobiography of his early career in biology entitled My Association with William Morton Wheeler. In this work, he describes vividly the reasons why he became a biologist, his studies and experiences at Rice Institute and his work at Harvard's Bussey Institute. Also included with George Wheeler's records is the memory book that he maintained during his stay at Rice Institute. In this scrapbook, Wheeler kept photographs of the biology staff, field trips and the campus, as well as newspaper and magazine clippings and memorabilia from social events.

ANT TAXONOMY
GEORGE CARLOS WHEELER, Doctor of Science in Applied Biology, Special Field, Economic Entomology. Thesis: "The Larvae of the Subfamily Dolichoderinae and Formicinae, with a General Consideration of the Developmental Stages of Ants."

PUBLICATIONS

 * [[Media:Deyrup 1989.pdf|Deyrup, M.; Johnson, C.; Wheeler, G. C.; Wheeler, J. 1989. A preliminary list of the ants of Florida. Fla. Entomol. 72: 91-101 PDF]]


 * [[Media:Wheeler GC 1928.pdf|Wheeler, G. C. 1928. The larva of Leptanilla (Hym.: Formicidae). Psyche (Camb.) 35: 85-91 PDF]]


 * [[Media:Wheeler G C 1935.pdf|Wheeler, G. C. 1935. The larva of Allomerus (Hym.: Formicidae). Psyche (Camb.) 42: 92-98 PDF]]


 * [[Media:Wheeler 1938.pdf|Wheeler, G. C. 1938. Are ant larvae apodous? Psyche (Cambridge) 45:139-145. [1938-12]Stamp date in MCZ library: 1939.03.01. PDF]]


 * [[Media:Wheeler 1943.pdf|Wheeler, G. C. 1943. The larvae of the army ants. Ann. Entomol. Soc. Am. 36: 319-332 PDF]]


 * [[Media:Wheeler 1949.pdf|Wheeler, G. C. 1949 [1948]. The larvae of the fungus-growing ants. Am. Midl. Nat. 40: 664-689 PDF]]


 * [[Media:Wheeler 1950.pdf|Wheeler, G. C. 1950. Ant larvae of the subfamily Cerapachyinae. Psyche (Camb.) 57: 102-113 PDF]]


 * [[Media:Wheeler & Wheeler 1930.pdf|Wheeler, G. C.; Wheeler, E. W. 1930. Two new ants from Java. Psyche (Camb.) 37: 193-201 PDF]]


 * [[Media:Wheeler & Wheeler 1934.pdf|Wheeler, G. C.; Wheeler, E. W. 1934. New forms of Aphaenogaster treatae Forel from the southern United States (Hym.: Formicidae). Psyche (Camb.) 41: 6-12 PDF]]


 * [[Media:Wheeler & Wheeler 1944.pdf|Wheeler, G. C.; Wheeler, E. W. 1944. The ants of North Dakota. N. D. Hist. Q. 11: 231-271 PDF]]


 * [[Media:Wheeler & Wheeler 1951.pdf|Wheeler, G. C.; Wheeler, J. 1951. The ant larvae of the subfamily Dolichoderinae. Proc. Entomol. Soc. Wash. 53: 169-210 PDF]]


 * [[Media:Wheeler & Wheeler 1952a.pdf|Wheeler, G. C.; Wheeler, J. 1952a. The ant larvae of the subfamily Ponerinae - Part I. Am. Midl. Nat. 48: 111-144 PDF]]


 * [[Media:Wheeler & Wheeler 1952b.pdf|Wheeler, G. C.; Wheeler, J. 1952b. The ant larvae of the myrmicine tribe Crematogastrini. J. Wash. Acad. Sci. 42: 248-262 PDF]]


 * [[Media:Wheeler & Wheeler 1952c.pdf|Wheeler, G. C.; Wheeler, J. 1952c. The ant larvae of the subfamily Ponerinae - Part II. Am. Midl. Nat. 48: 604-672 PDF]]


 * [[Media:Wheeler & Wheeler 1953a.pdf|Wheeler, G. C.; Wheeler, J. 1953a [1952]. The ant larvae of the myrmicine tribe Myrmicini. Psyche (Camb.) 59: 105-125 PDF]]


 * [[Media:Wheeler & Wheeler 1953b.pdf|Wheeler, G. C.; Wheeler, J. 1953b. The ant larvae of the myrmicine tribe Pheidolini (Hymenoptera, Formicidae). Proc. Entomol. Soc. Wash. 55: 49-84 PDF]]


 * [[Media:Wheeler & Wheeler 1953c.pdf|Wheeler, G. C.; Wheeler, J. 1953c. The ant larvae of the subfamily Formicinae. Ann. Entomol. Soc. Am. 46: 126-171 PDF]]


 * [[Media:Wheeler & Wheeler 1953d.pdf|Wheeler, G. C.; Wheeler, J. 1953d. The ant larvae of the myrmicine tribes Melissotarsini, Metaponini, Myrmicariini, and Cardiocondylini. J. Wash. Acad. Sci. 43: 185-189 PDF]]


 * [[Media:Wheeler & Wheeler 1953e.pdf|Wheeler, G. C.; Wheeler, J. 1953e. The ant larvae of the subfamily Formicinae. Part II. Ann. Entomol. Soc. Am. 46: 175-217 PDF]]


 * [[Media:Wheeler & Wheeler 1954a.pdf|Wheeler, G. C.; Wheeler, J. 1954a [1953]. The ant larvae of the myrmicine tribe Pheidologetini. Psyche (Camb.) 60: 129-147 PDF]]


 * [[Media:Wheeler & Wheeler 1954b.pdf|Wheeler, G. C.; Wheeler, J. 1954b. The ant larvae of the myrmicine tribes Cataulacini and Cephalotini. J. Wash. Acad. Sci. 44: 149-157 PDF]]


 * [[Media:Wheeler & Wheeler 1954c.pdf|Wheeler, G. C.; Wheeler, J. 1954c. The ant larvae of the myrmicine tribe Myrmecinini (Hymenoptera). Proc. Entomol. Soc. Wash. 56: 126-138 PDF]]


 * [[Media:Wheeler & Wheeler 1954d.pdf|Wheeler, G. C.; Wheeler, J. 1954d. The ant larvae of the myrmicine tribes Meranoplini, Ochetomyrmicini and Tetramoriini. Am. Midl. Nat. 52: 443-452 PDF]]


 * [[Media:Wheeler & Wheeler 1955a.pdf|Wheeler, G. C.; Wheeler, J. 1955a [1954]. The ant larvae of the myrmicine tribes Basicerotini and Dacetini. Psyche (Camb.) 61: 111-145 PDF]]


 * [[Media:Wheeler & Wheeler 1955b.pdf|Wheeler, G. C.; Wheeler, J. 1955b. The ant larvae of the myrmicine tribe Leptothoracini. Ann. Entomol. Soc. Am. 48: 17-29 PDF]]


 * [[Media:Wheeler and Wheeler 1955c.pdf|Wheeler, G. C.; Wheeler, J. 1955c. The ant larvae of the myrmicine tribe Solenopsidini. Am. Midl. Nat. 54: 119-141 PDF]]


 * [[Media:Wheeler & Wheeler 1956.pdf|Wheeler, G. C.; Wheeler, J. 1956. The ant larvae of the subfamily Pseudomyrmecinae (Hymenoptera: Formicidae). Ann. Entomol. Soc. Am. 49: 374-398 PDF]]


 * [[Media:Wheeler & Wheeler 1957a.pdf|Wheeler, G. C.; Wheeler, J. 1957a. The larva of the ant genus Dacetinops Brown and Wilson. Breviora 78: 1-4 PDF]]


 * [[Media:Wheeler & Wheeler 1957b.pdf|Wheeler, G. C.; Wheeler, J. 1957b. The larva of Simopelta (Hymenoptera: Formicidae). Proc. Entomol. Soc. Wash. 59: 191-194 PDF]]


 * [[Media:Wheeler & Wheeler 1957c.pdf|Wheeler, G. C.; Wheeler, J. 1957c [1956]. Veromessor lobognathus in North Dakota (Hymenoptera: Formicidae). Psyche (Camb.) 63: 140-145 PDF]]


 * [[Media:Wheeler & Wheeler 1959b.pdf|Wheeler, G. C.; Wheeler, J. 1959b. The larva of Paramyrmica (Hymenoptera: Formicidae). J. Tenn. Acad. Sci. 34: 219-220 PDF]]


 * [[Media:Wheeler & Wheeler 1960a.pdf|Wheeler, G. C.; Wheeler, J. 1960a. The ant larvae of the subfamily Myrmicinae. Ann. Entomol. Soc. Am. 53: 98-110 PDF]]


 * [[Media:Wheeler & Wheeler 1960b.pdf|Wheeler, G. C.; Wheeler, J. 1960b. Supplementary studies on the larvae of the Myrmicinae (Hymenoptera: Formicidae). Proc. Entomol. Soc. Wash. 62: 1-32 PDF]]


 * [[Media:Wheeler & Wheeler 1960c.pdf|Wheeler, G.C.& J.Wheeler 1960c. Techniques for the Study of Ant Larvae. Psyche 67 (4):87-94 PDF]]


 * Wheeler, G. C.; Wheeler, J. 1963. The ants of North Dakota. Grand Forks, North Dakota: University of North Dakota Press, viii + 326 pp.


 * [[Media:Wheeler & Wheeler 1964a.pdf|Wheeler, G. C.; Wheeler, J. 1964a. The ant larvae of the subfamily Cerapachyinae: supplement. Proc. Entomol. Soc. Wash. 66: 65-71 PDF]]


 * [[Media:Wheeler & Wheeler 1964b.pdf|Wheeler, G. C.; Wheeler, J. 1964b. The ant larvae of the subfamily Ponerinae: supplement. Ann. Entomol. Soc. Am. 57: 443-462 PDF]]


 * [[Media:Wheeler & Wheeler 1964c.pdf|Wheeler, G. C.; Wheeler, J. 1964c. The ant larvae of the subfamily Dorylinae: supplement. Proc. Entomol. Soc. Wash. 66: 129-137 PDF]]


 * [[Media:Wheeler & Wheeler 1965b.pdf|Wheeler, G. C.; Wheeler, J. 1965b. The ant larvae of the subfamily Leptanillinae (Hymenoptera, Formicidae). Psyche (Camb.) 72: 24-34 PDF]]


 * [[Media:Wheeler & Wheeler 1966.pdf|Wheeler, G. C.; Wheeler, J. 1966. Ant larva of the subfamily Dolichoderinae: supplement. Ann. Entomol. Soc. Am. 59: 726-732 PDF]]


 * [[Media:Wheeler & Wheeler 1968a.pdf|Wheeler, G. C.; Wheeler, J. 1968a. The ant larvae of the subfamily Formicinae (Hymenoptera: Formicidae): supplement. Ann. Entomol. Soc. Am. 61: 205-222 PDF]]


 * [[Media:Wheeler & Wheeler 1969.pdf|Wheeler, G. C.; Wheeler, J. 1969. The larva of Acanthognathus (Hymenoptera: Formicidae). Psyche (Camb.) 76: 110-113 PDF]]


 * [[Media:Wheeler & Wheeler 1970b.pdf|Wheeler, G. C.; Wheeler, J. 1970b. Ant larvae of the subfamily Formicinae: second supplement. Ann. Entomol. Soc. Am. 63: 648-656 PDF]]


 * [[Media:Wheeler & Wheeler 1971a.pdf|Wheeler, G. C.; Wheeler, J. 1971a [1970]. The larva of Apomyrma (Hymenoptera: Formicidae). Psyche (Camb.) 77: 276-279 PDF]]


 * [[Media:Wheeler & Wheeler 1971b.pdf|Wheeler, G. C.; Wheeler, J. 1971b. Ant larvae of the subfamily Ponerinae: second supplement. Ann. Entomol. Soc. Am. 6 64: 1197-1217 PDF]]


 * [[Media:Wheeler & Wheeler 1971c.pdf|Wheeler, G. C.; Wheeler, J. 1971c. The larvae of the ant genus Bothroponera (Hymenoptera: Formicidae). Proc. Entomol. Soc. Wash. 73: 386-394 PDF]]


 * [[Media:Wheeler & Wheeler 1971d.pdf|Wheeler, G. C.; Wheeler, J. 1971d. Ant larvae of the subfamily Myrmeciinae (Hymenoptera: Formicidae). Pan-Pac. Entomol. 47: 245-256 PDF]]


 * [[Media:Wheeler & Wheeler 1972a.pdf|Wheeler, G. C.; Wheeler, J. 1972a. The subfamilies of Formicidae. Proc. Entomol. Soc. Wash. 74: 35-45 PDF]]


 * [[Media:Wheeler & Wheeler 1972b.pdf|Wheeler, G. C.; Wheeler, J. 1972b. Ant larvae of the subfamily Myrmicinae: second supplement on the tribes Myrmicini and Pheidolini. J. Ga. Entomol. Soc. 7: 233-246 PDF]]


 * [[Media:Wheeler & Wheeler 1973a.pdf|Wheeler, G. C.; Wheeler, J. 1973a. The ant larvae of six tribes: second supplement (Hymenoptera: Formicidae: Myrmicinae). J. Ga. Entomol. Soc. 8: 27-39 PDF]]


 * [[Media:Wheeler & Wheeler 1973b.pdf|Wheeler, G. C.; Wheeler, J. 1973b. Ant larvae of four tribes: second supplement (Hymenoptera: Formicidae: Myrmicinae). Psyche (Camb.) 80: 70-82 PDF]]


 * [[Media:Wheeler & Wheeler 1973c.pdf|Wheeler, G. C.; Wheeler, J. 1973c. The ant larvae of the tribes Basicerotini and Dacetini: second supplement (Hymenoptera: Formicidae: Myrmicinae). Pan-Pac. Entomol. 49: 207-214 PDF]]


 * [[Media:Wheeler & Wheeler 1973.pdf|Wheeler, G. C.; Wheeler, J. 1973d. Supplementary studies on ant larvae: Cerapachyinae, Pseudomyrmecinae and Myrmicinae. Psyche (Camb.) 80: 204-211 PDF]]


 * Wheeler, G. C.; Wheeler, J. 1973e. Ants of Deep Canyon. Riverside, Calif.: University of California, xiii + 162 pp.


 * [[Media:Wheeler & Wheeler 1974a.pdf|Wheeler, G. C.; Wheeler, J. 1974a. Ant larvae of the subfamily Formicinae: third supplement. J. Ga. Entomol. Soc. 9: 59-64 PDF]]


 * [[Media:Wheeler & Wheeler 1974b.pdf|Wheeler, G. C.; Wheeler, J. 1974b [1973]. Ant larvae of the subfamily Dolichoderinae: second supplement (Hymenoptera: Formicidae). Pan-Pac. Entomol. 49: 396-401 PDF]]


 * [[Media:Wheeler & Wheeler 1974c.pdf|Wheeler, G. C.; Wheeler, J. 1974c. Ant larvae of the subfamily Dorylinae: second supplement (Hymenoptera: Formicidae). J. Kans. Entomol. Soc. 47: 166-172 PDF]]


 * [[Media:Wheeler & Wheeler 1974d.pdf|Wheeler, G. C.; Wheeler, J. 1974d. Ant larvae of the myrmicine tribe Attini: second supplement (Hymenoptera: Formicidae). Proc. Entomol. Soc. Wash. 76: 76-81 PDF]]


 * [[Media:Wheeler & Wheeler 1974.pdf|Wheeler, G. C.; Wheeler, J. 1974e. Supplementary studies on ant larvae: Teratomyrmex. Psyche (Camb.) 81: 38-41 PDF]]


 * [[Media:Wheeler & Wheeler 1974f.pdf|Wheeler, G. C.; Wheeler, J. 1974f. Supplementary studies on ant larvae: Simopone and Turneria. J. N. Y. Entomol. Soc. 82: 103-105 PDF]]


 * [[Media:Wheeler & Wheeler 1974g.pdf|Wheeler, G. C.; Wheeler, J. 1974g. Ant larvae of the subfamily Ponerinae: third supplement (Hymenoptera: Formicidae). Proc. Entomol. Soc. Wash. 76: 278-281 PDF]]


 * [[Media:Wheeler & Wheeler 1976a.pdf|Wheeler, G. C.; Wheeler, J. 1976a. Supplementary studies on ant larvae: Ponerinae. Trans. Am. Entomol. Soc. 102: 41-64 PDF]]


 * [[Media:Wheeler & Wheeler 1976b.pdf|Wheeler, G. C.; Wheeler, J. 1976b. Ant larvae: review and synthesis. Mem. Entomol. Soc. Wash. 7: 1-108 PDF]]


 * [[Media:Wheeler & Wheeler 1977a.pdf|Wheeler, G. C.; Wheeler, J. 1977a. Supplementary studies on ant larvae: Myrmicinae. Trans. Am. Entomol. Soc. 103: 581-602 PDF]]


 * [[Media:Wheeler & Wheeler 1977b.pdf|Wheeler, G. C.; Wheeler, J. 1977b. North Dakota ants updated. Reno, Nevada: published by the authors, 27 pp. PDF]]


 * [[Media:Wheeler & Wheeler 1978.pdf|Wheeler, G.C.; Wheeler, J. 1978. Mountain ants of Nevada. Great Basin Naturalist 38: 379-396 PDF]]


 * [[Media:Wheeler & Wheeler 1980.pdf|Wheeler, G. C.; Wheeler, J. 1980. Supplementary studies on ant larvae: Ponerinae, Myrmicinae and Formicinae. Trans. Am. Entomol. Soc. 106: 527-545 PDF]]


 * [[Media:Wheeler & Wheeler 1982.pdf|Wheeler, G. C.; Wheeler, J. 1982b. Supplementary studies on ant larvae: Formicinae (Hymenoptera: Formicidae). Psyche (Camb.) 89: 175-181 PDF]]


 * [[Media:Wheeler & Wheeler 1983a.pdf|Wheeler, G. C.; Wheeler, J. 1983a. Supplementary studies on ant larvae: Myrmicinae. Trans. Am. Entomol. Soc. 108: 601-610 PDF]]


 * [[Media:Wheeler & Wheeler 1984a.pdf|Wheeler, G. C.; Wheeler, J. 1984a. The larvae of the army ants (Hymenoptera: Formicidae): a revision. J. Kans. Entomol. Soc. 57: 263-275 PDF]]


 * Wheeler, G. C.; Wheeler, J. 1984b. Myrmecological orthoepy and onomatology. (2nd edition.). Grand Forks, North Dakota: published by the authors, 20 pp


 * [[Media:Wheeler & Wheeler 1985.pdf|Wheeler, G. C. and J. Wheeler. 1985a. A checklist of Texas ants. The Prairie Naturalist 17(2): 49-64. PDF]]


 * [[Media:Wheeler & Wheeler 1985b.pdf|Wheeler, G. C.; Wheeler, J. 1985b. A simplified conspectus of the Formicidae. Trans. Am. Entomol. Soc. 111: 255-264 PDF]]


 * [[Media:Wheeler & Wheeler 1986a.pdf|Wheeler, G. C.; Wheeler, J. 1986a [1985]. The larva of Dinoponera (Hymenoptera: Formicidae: Ponerinae). Psyche (Camb.) 92: 387-39 PDF]]


 * [[Media:Wheeler & Wheeler 1985.pdf|Wheeler, G. C.; Wheeler, J. 1986b [1985]. The larva of Proatta (Hymenoptera: Formicidae). Psyche (Camb.) 92: 447-450 PDF]]


 * [[Media:Wheeler & Wheeler 1986c.pdf|Wheeler, G. C.; Wheeler, J. 1986c. Supplementary studies of ant larvae: Ponerinae. Trans. Am. Entomol. Soc. 112: 85-94 PDF]]


 * [[Media:Wheeler & Wheeler 1986d.pdf|Wheeler, G. C.; Wheeler, J. 1986d. Supplementary studies on ant larvae: Formicinae (Hymenoptera: Formicidae). J. N. Y. Entomol. Soc. 94: 331-341 PDF]]


 * [[Media:Wheeler & Wheeler 1986f.pdf|Wheeler, G. C.; Wheeler, J. 1986f. Supplementary studies on ant larvae: Myrmicinae (Hymenoptera: Formicidae). J. N. Y. Entomol. Soc. 94: 489-499 PDF]]


 * [[Media:Wheeler & Wheeler 1986g.pdf| Wheeler, G. C.; Wheeler, J. 1986g. The ants of Nevada. Los Angeles: Natural History Museum of Los Angeles County, vii + 138 pp. PDF]]


 * [[Media:Wheeler & Wheeler 1986.pdf|Wheeler, G. C.; Wheeler, J. 1987a [1986]. Young larvae of Eciton (Hymenoptera: Formicidae: Dorylinae). Psyche (Camb.) 93: 341-349 PDF]]


 * [[Media:Wheeler & Wheeler 1988b.pdf|Wheeler, G. C.; Wheeler, J. 1988b [1987]. Young larvae of Veromessor pergandei (Hymenoptera: Formicidae: Myrmicinae). Psyche (Camb.) 94: 303-307 PDF]]


 * [[Media:Wheeler & Wheeler 1988c.pdf|Wheeler, G. C.; Wheeler, J. 1988c. The larva of Notostigma (Hymenoptera: Formicidae: Formicinae). J. N. Y. Entomol. Soc. 96: 355-358 PDF]]


 * [[Media:Wheeler & Wheeler 1989a.pdf|Wheeler, G. C.; Wheeler, J. 1989a [1988]. Notes on ant larvae: Myrmicinae. Trans. Am. Entomol. Soc. 114: 319-327 PDF]]


 * [[Media:Wheeler & Wheeler 1989b.pdf|Wheeler, G. C.; Wheeler, J. 1989b. Notes on ant larvae: Ponerinae. J. N. Y. Entomol. Soc. 97: 50-55 PDF]]


 * [[Media:Wheeler & Wheeler 1989.pdf|Wheeler, G. C.; Wheeler, J. 1989c [1988]. The larva of Leptanilla japonica, with notes on the genus (Hymenoptera: Formicidae: Leptanillinae). Psyche (Camb.) 95: 185-189 PDF]]


 * [[Media:Wheeler & Wheeler 1988.pdf|Wheeler, G. C.; Wheeler, J. 1988d. A checklist of the ants of Montana. Psyche (Cambridge) 95:101-114. [1988-12-20] PDF]]


 * [[Media:Wheeler & Wheeler 1990a.pdf|Wheeler, G. C.; Wheeler, J. 1990a [1989]. Notes on ant larvae. Trans. Am. Entomol. Soc. 115: 457-473 PDF]]


 * [[Media:Wheeler & Wheeler 1990b.pdf|Wheeler, G. C.; Wheeler, J. 1990b. Larvae of the formicine ant genus Polyrhachis. Trans. Am. Entomol. Soc. 116: 753-767 PDF]]


 * [[Media:Wheeler & Wheeler 1991a.pdf|Wheeler, G. C.; Wheeler, J. 1991a. The larva of Blepharidatta (Hymenoptera: Formicidae). J. N. Y. Entomol. Soc. 99: 132-137 PDF]]


 * [[Media:Wheeler & Wheeler 1991b.pdf|Wheeler, G. C.; Wheeler, J. 1991b. Instars of three ant species. Psyche (Camb.) 98: 89-99 PDF]]


 * [[Media:Wheeler Wheeler Kannowski 1994.pdf|Wheeler, G. C.; Wheeler, J.; Kannowski, P. B. 1994. Checkist of ants of Michigan (Hymenoptera: Formicidae). Great Lakes Entomol. 26: 297-310 PDF]]


 * Wheeler, G. C.; Wheeler, J.; Taylor, R. W. 1980. The larval and egg stages of the primitive ant Nothomyrmecia macrops Clark (Hymenoptera: Formicidae). J. Aust. Entomol. Soc. 19: 131-137


 * [[Media:Wilson et al 1956.pdf|Wilson, E. O.; Eisner, T.; Wheeler, G. C.; Wheeler, J. 1956. Aneuretus simoni Emery, a major link in ant evolution. Bull. Mus. Comp. Zool. 115: 81-99 PDF]]