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Genus Gryllus

field crickets

link to keys Key to genera of field crickets (Gryllinae).

Species of the genus Gryllus (field crickets) are the most-studied of North American singing insects. They are large, easy to rear and handle, and diverse in their acoustic behavior, ecology, and life cycles. Their early taxonomic history is instructive. Many species were described in the 1800s, but when Rehn and Hebard (1915) intensively analyzed 1,500 pinned Western Hemisphere specimens, they concluded that all native American Gryllus belonged to a single "exceedingly plastic" species, Gryllus assimilis. This conclusion held for more than 30 years, until B. B. Fulton (1952) studied, in the field and in captivity, living Gryllus in North Carolina. He showed there were four reproductively isolated populations that differed in their calling songs, habitats, and seasonal life cycles. Because he found no defining morphological differences, Fulton did not give scientific names to the species he had discovered. R. D. Alexander (1957) extended Fulton’s studies of Gryllus to the Midwest, found a fifth species, and showed that most species pairs were separable by morphological characters, at least in one sex. He assigned scientific names, thus confronting museum curators with the fact that their trays of Gryllus assimilis probably contained mixtures of species none of which was likely to be G. assimilis. More importantly, differentiating and naming species of North American Gryllus made them inviting subjects for studies of evolution, behavior, and physiology.

Since 1957, six more Gryllus species have been recognized in the eastern United States and 5 additional ones have been found in the West. David Weissman is currently working on a revision of Gryllus from the western and central states that will probably more than double the number of recognized species.

Studies of calling songs and life histories are still the predominant means of recognizing new species of Gryllus. Once a new species is recognized, morphological differences may be easy or difficult to find.

Seasonal life cycles

Species of Gryllus from the same geographic locality may have very different seasonal life histories. In fact, the two most abundant field crickets in the Northeast are separable chiefly by their life histories. Gryllus veletis and G. pennsylvanicus (spring and fall field crickets) do not differ in song or habitat and differ morphologically only in the average length of the ovipositor relative to the body length. However, G. veletis overwinters as mid-sized juveniles and matures in spring, whereas G. pennsylvanicus overwinters as eggs and matures in fall. The two species co-occur as adults, in very small numbers, only briefly in midsummer. Thus, except for the occasional specimen collected during the period of overlap, the date of collection is sufficient to distinguish adults of the two species.

G. veletis and G. pennsylvanicus, like many other Gryllus, have a single annual generation. Some species have two discrete generations per year (e.g., G. rubens) and others have generations that overlap (e.g. G. assimilis). G. firmus in Gainesville, Florida, has remarkably varied responses to seasons. It overwinters in all stages except small juveniles, and the eggs of a single female may hatch over a five-month period and the resulting juveniles may mature over a nine month period!

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Representative life cycles Asynchronous development Phylogeny


The evolutionary branching sequence (phylogeny) that produced the species of North American field crickets is important to those wishing to understand the evolution of the differences among the species. For example, the similarity of G. pennsylvanicus and G. veletis, except in their seasonal life cycles, led to the assumption that the two were sister species and to a theory of how the ancestral species might have produced the two species sympatrically by allochronic speciation (Alexander and Bigelow 1960). Subsequently, studies of chromosomes, allozymes, and mitochondrial DNA indicated that the split between egg and juvenile-overwintering species occurred early in the phylogeny of North American Gryllus and that G. pennsylvanicus and G. veletis were on different branches of the Gryllus family tree.

Huang et al. (2000) reconstructed the phylogeny of ten species of North American Gryllus based on the DNA sequences of two mitochondrial genes. Trios of closely related species were firmus-ovisopis-pennsylvanicus, texensis-rubens-lineaticeps, and fultoni-integer-veletis. G. assimilis was not closely related to any of the other species but it may form a monophyletic group with fultoni-integer-veletis.


Alexander RD. 1957. The taxonomy of the field crickets of the eastern United States (Orthptera: Gryllidae: Acheta). Ann. Entomol. Soc. Am. 50: 584-602.

Alexander RD, Bigelow RS. 1960. Allochronic speciation in field crickets, and a new species, Acheta veletis. Evolution 14: 334-346.

Alexander RD, Meral GH. 1967. Seasonal and daily chirping cycles in the northern spring and fall field crickets Gryllus veletis and G. pennsylvanicus. Ohio J. Sci. 67: 200-209.

Alexander RD, Walker TJ. 1962. Two introduced field crickets new to eastern United States (Orthoptera, Gryllidae). Ann. Entomol. Soc. Am. 55(1): 90-94. [Gryllus assimilis, Velarifictorus micado]

Backus VL, Cade WH. 1986. Sperm competition in the field cricket Gryllus integer (Orthoptera: Gryllidae). Fla. Entomol. 69: 722-728. [=Gryllus texensis]

Bertram SM. 2000. The influence of age and size on temporal mate signalling behaviour. Anim. Behav. 60: 333-339. [Gryllus texensis]

Bigelow RS. 1958. Evolution in the field cricket Acheta assimilis Fab. Can. J. Zool. 36: 139-151. [=Gryllus spp.]

Bigelow RS. 1960a. Developmental rates and diapause in Acheta pennsylvanicus (Burmeister) and Acheta veletis Alexander and Bigelow (Orthoptera: Gryllidae). Can. J. Zool. 38: 973-988.

Bigelow RS. 1960b. interspecific hybrids and speciation in the genus Acheta (Orthoptera Gryllidae). Can. J. Zool. 38: 509-524. [=Gryllus spp.]

Cade WH. 1979. Effect of male-deprivation on female phonotaxis in field crickets (Orthoptera, Gryllidae; Gryllus). Can. Entomol. 111: 741-744. [Gryllus veletis and G. texensis]

Cade WH. 1981a. Field cricket spacing, and the phonotaxis of crickets and parasitoid flies to clumped and isolated cricket songs. Z. Tierpsychol. 55: 365-375. [Gryllus veletis and G. texensis]

Cade WH. 1981b. Alternative male strategies: Genetic differences in crickets (Gryllus integer). Science 212: 563-564. [=Gryllus texensis]

Cade WH. 1984. Effects of fly parasitoids on nightly calling duration in field crickets (Gryllus integer). Can. J. Zool. 62: 226-228. [=Gryllus texensis]

Cade WH. 1989. Nightly and hourly rates of attraction of flying field crickets, Gryllus integer, to conspecific song. Can. J. Zool. 67: 2540-2542. [=Gryllus texensis]

Cade WH. 1991. Interspecific and intraspecific variation in nightly calling duration in field crickets, Gryllus integer and Gryllus rubens (Orthoptera: Gryllidae). J. insect Behav. 4: 185-194. ["Gryllus integer"= G. texensis]

Cade WH, Cade ES. 1992. Male mating success, calling and searching behaviour at high and low densities in the field cricket, Gryllus integer. Anim. Behav. 43: 49-56. [=Gryllus texensis]

Cade WH, Ciceran M, Murray AM. 1996. Temporal patterns of parasitoid fly (Ormia ochracea) attraction to field cricket song (Gryllus integer). Can. J. Zool. 74: 393-395. [=Gryllus texensis]

Cade WH, Otte D. 2000. Gryllus texensis n. sp.: a widely studied field cricket (Orthoptera; Gryllidae) from the southern United States. Trans. Am. Entomol. Soc. 126: 117-123.

Cade WH, Tyshenko MG. 1990. Geographic variation in hybrid fertility in the field crickets Gryllus integer, Gryllus rubens, and Gryllus sp. Can. J. Zool. 68: 2697-2700. ["Gryllus sp."= G. texensis]

Cade WH, Wyatt DR. 1984. Factors affecting calling behavior in field crickets, Teleogryllus and Gryllus: Age, weight, density and parasites. Behaviour 88: 61-75. [G. texensis, G. pennsylvanicus, G. veletis]

Carriere Y, Roff DA. 1995. The evolution of offspring size and number: A test of the Smith-Fretwell model in three species of crickets. Oecologia 102: 389-396. [Gryllus firmus, G. veletis, G. pennsylvanicus]

Carriere Y, Simons AM, Roff DA. 1996. The effect of the timing of post-diapause egg development on survival, growth, and body size in Gryllus pennsylvanicus. Oikos 75: 463-470.

Ciceran M, Murray AM, Rowell G. 1994. Natural variation in the temporal patterning of calling song structure in the field cricket Gryllus pennsylvanicus: Effects of temperature, age, mass, time of day, and nearest neighbour. Can. J. Zool. 72: 38-42.

Dixon KA, Cade WH. 1986. Some factors influencing male-male aggression in the field cricket Gryllus integer (time of day, age, weight and sexual maturity). Anim. Behav. 34: 340-346. [=Gryllus texensis]

Doherty JA, Callos JD. 1991. Acoustic communication in the trilling field cricket, Gryllus rubens (Orthoptera: Gryllidae). J. insect Behav. 4: 67-82.

Doherty JA, Storz MM. 1992. Calling song and selective phonotaxis in the field crickets, Gryllus firmus and Gryllus pennsylvanicus (Orthoptera: Gryllidae). J. insect Behav. 5: 555-569.

Farris HE, Forrest TG, Hoy RR. 1998. The effect of ultrasound on the attractiveness of acoustic mating signals. Physiol. Entomol. 23: 322-328. [Gryllus rubens]

Fitzpatrick MJ, Gray DA. 2001. Divergence between the courtship songs of the field crickets Gryllus texensis and Gryllus rubens (Orthoptera: Gryllidae). Ethology 107: 1075-1085.

French BW, Cade WH. 1987. The timing of calling, movement, and mating in the field crickets Gryllus veletis, Gryllus pennsylvanicus, and Gryllus integer. Behav. Ecol. Sociobiol. 21: 157-162.["G. integer" = G. texensis]

French BW, Cade WH. 1989. Sexual selection at varying population densities in male field crickets, Gryllus veletis and Gryllus pennsylvanicus. J. insect Behav. 2: 105-122.

Fulton BB. 1952. Speciation in the field cricket. Evolution 6:283-95.

Giordano R, Jackson JJ, Robertson HM. 1997. The role of Wolbachia bacteria in reproductive incompatibilities and hybrid zones of Diabrotica beetles and Gryllus crickets. Proc. Natl. Acad. Sci. USA 94: 11439-11444.

Gray DA, Cade WH. 1999. Quantitative genetics of sexual selection in the field cricket, Gryllus integer. Evolution 53: 848-854. [=Gryllus texensis]

Gray DA, Cade WH. 1999. Sex, death and genetic variation: Natural and sexual selection on cricket song. Proc. R. Soc. Biol. Sci. Ser. B 266: 707-709. [Gryllus texensis]

Gray DA, Cade WH. 2000. Sexual selection and speciation in field crickets. PNAS 97: 14449-14454. [Gryllus texensis and G. rubens]

Gray DA, Walker TJ, Conley BE, Cade WH. 2001. A morphological means of distinguishing females of the cryptic field cricket species, Gryllus rubens and G. texensis (Orthoptera: Gryllidae). Fla. Entomol. 84: 314-315.

Gurney AB. 1950. Entomology--The Linnaean subgeneric names of Gryllus (Orthoptera). J. Wash. Acad. Sci. 40: 409-413.

Harrison RG. 1979. Flight polymorphism in the field cricket Gryllus pennsylvanicus. Oecologia 40: 125-132.

Harrison RG. 1979. Speciation in North American field crickets: Evidence from electrophoretic comparisons. Evolution 33: 1009-1023.

Harrison RG, Arnold J. 1982. A narrow hybrid zone between closely related cricket species. Evolution 36: 535-552. [Gryllus pennsylvanicus and G. firmus]

Harrison RG. 1983. Barriers to gene exchange between closely related cricket species: 1. Laboratory hybridization studies. Evolution 37: 245-251. [Gryllus pennsylvanicus and G. firmus]

Harrison RG. 1985. Barriers to gene exchange between closely related cricket species: 2. Life cycle variation and temporal isolation. Evolution 39: 244-259. [Gryllus pennsylvanicus and G. firmus]

Harrison RG. 1986. Pattern and process in a narrow hybrid zone. Heredity 56: 337-350. [Gryllus pennsylvanicus and G. firmus]

Harrison RG, Rand DM, Wheeler WC. 1987. Mitochondrial DNA variation in field crickets across a narrow hybrid zone. Mol. Biol. Evol. 4: 144-158. [Gryllus pennsylvanicus and G. firmus]

Harrison RG, Bogdanowicz SM. 1995. Mitochondrial DNA phylogeny of North American field crickets: perspectives on the evolution of life cycles, songs, and habitat associations. J. Evol. Biol. 8: 209-232.

Harrison RG, Bogdanowicz SM. 1997. Patterns of variation and linkage disequilibrium in a field cricket hybrid zone. Evolution 51: 493-505.

Hedrick AV. 1986. Female preferences for male calling bout duration in a field cricket (Gryllus integer). Behav. Ecol. Sociobiol. 19: 73-77.

Hedrick AV. 1988. Female choice and the heritability of attractive male traits: An empirical study. Am. Nat. 132: 267-276. [Gryllus integer]

Hedrick A, Weber T. 1998. Variance in female responses to the fine structure of male song in the field cricket, Gryllus integer. Behav. Ecol. 9: 582-591.

Hedrick AV. 2000. Crickets with extravagant mating songs compensate for predation risk with extra caution. Proc. R. Soc. Biol. Sci. Ser. B 267: 671-675. [Gryllus integer]

Hoback WW, Wagner WE, Jr. 1997. The energetic cost of calling in the variable field cricket, Gryllus lineaticeps. Physiol. Entomol. 22: 286-290.

Holtmeier CL, Zera AJ. 1993. Differential mating success of male wing morphs of the cricket, Gryllus rubens. Am. Midl. Nat. 129: 223-233.

Huang Y, Orti G, Sutherlin M, Siegel-Causey D, Duhachek A, Zera AJ. 2000. Phylogenetic relationships of North American field crickets inferred from mitochondrial DNA data. Mol. Phylogenet. Evol. 17: 48-57.

Ibrahim R, Walker TJ. 1980. Diapause and nondiapause eggs laid daily by individual Gryllus firmus females (Orthoptera: Gryllidae).  Fla. Entomol. 63(4): 510-512.

Jang, Y, Gerhardt, HC. 2005. Divergence in the callling songs between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae). J. Evol. Biol. 19: 459–472.

Jang Y, Gerhardt, HC. 2006. Divergence in female calling song discrimination between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae). Behav. Ecol. Sociobiol.
Online First, DOi 10.1007/s00265-005-0151-3, URL

Lickman K, Murray AM, Cade WH. 1998. Effect of mating on female phonotactic response in Gryllus integer (Orthoptera: Gryllidae). Can. J. Zool. 76: 1263-1268. [=Gryllus texensis]

Mole S, Zera AJ. 1993. Differential allocation of resources underlies the dispersal-reproduction trade-off in the wing-dimorphic cricket, Gryllus rubens. Oecologia 93: 121-127.

Murray AM, Cade WH. 1995. Differences in age structure among field cricket populations (Orthoptera: Gryllidae): Possible influence of a sex-biased parasitoid. Can. J. Zool. 73: 1207-1213. [Gryllus veletis, G. pennsylvanicus, G. texensis]

Nickle DA, Walker TJ. 1974. A morphological key to field crickets of southeastern United States (Orthoptera: Gryllidae: Gryllus). Fla. Entomol. 57: 8-12.

Paul RC. 1977. Species-specific phonotaxis in Gryllus females (Note). Fla. Entomol. 60: 67-68.

Pires A, Hoy RR. 1992. Temperature coupling in cricket acoustic communication: i. Field and laboratory studies of temperature effects on calling song production and recognition in Gryllus firmus. J. Comp. Physiol. A. Sens. Neural. Behav. Physiol. 171: 69-78.

Pires A, Hoy RR. 1992. Temperature coupling in cricket acoustic communication: ii. Localization of temperature effects on song production and recognition networks in Gryllus firmus. J. Comp. Physiol. A. Sens. Neural. Behav. Physiol. 171: 79-92.

Prosser MR, Murray AM, Cade WH. 1997. The influence of female age on phonotaxis during single and multiple song presentations in the field cricket, Gryllus integer (Orthoptera: Gryllidae). J. insect Behav. 10: 437-449. [=G.texensis]

Rakshpal R. 1960. Sound-producing organs and mechanism of song production in field crickets of the genus Acheta fabricius (Orthoptera Gryllidae). Can. J. Zool. 38: 499-507. [=Gryllus spp.]

Rakshpal R. 1962a. The effect of cold on pre- and post-diapause eggs of Gryllus pennsylvanicus Burmeister (Orthoptera: Gryllidae). Proc. R. Entomol. Soc. Lond. 37: 117-120.

Rakshpal R. 1962b. Diapause in the eggs of Gryllus pennsylvanicus Burmeister (Orthoptera: Gryllidae). Can. J. Zool. 40: 179-194.

Rand DM, Harrison RG. 1989. Ecological genetics of a mosaic hybrid zone: Mitochondrial, nuclear, and reproductive differentiation of crickets by soil type. Evolution 43: 432-449. [Gryllus pennsylvanicus and G. firmus]

Randell RL, Kevan DKM. 1962. A cytological study of certain American species of Gryllus Linne (Orthoptera: Gryllidae) and their hybrids. Ann. Entomol. Soc. Que. 7: 48-60.

Reale D, Roff DA. 2002. Quantitative genetics of oviposition behaviour and interactions among oviposition traits in the sand cricket. Anim. Behav. 64: 397-406.

Rehn JAG, Hebard M. 1915. The genus Gryllus (Orthoptera) as found in America. Proc. Acad. Nat. Sci. 67: 293-322.

Roff A. 1990. Selection for changes in the incidence of wing dimorphism in Gryllus firmus. Heredity 65: 163-168.

Roff DA, Shannon P. 1993. Genetic and ontogenetic variation in behavior: its possible role in the maintenance of genetic variation in the wing dimorphism of Gryllus firmus. Heredity 71: 481-487.

Roff DA, Simons AM. 1997. The quantitative genetics of wing dimorphism under laboratory and 'field' conditions in the cricket Gryllus pennsylvanicus. Heredity 78: 235-240.

Ross CL, Harrison RG. 2002. A fine-scale spatial analysis of the mosaic hybrid zone between Gryllus firmus and Gryllus pennsylvanicus. Evolution 56: 2296-2312.

Rowel GA, Cade WH. 1993. Simulation of alternative male reproductive behavior: Calling and satellite behavior in field crickets. Ecol. Model. 65: 265-280. [G. texensis]

Smith CJ, Cade WH. 1987. Relative fertility in hybridization experiments using three song types of the field crickets Gryllus integer and Gryllus rubens. Can. J. Zool. 65: 2390-2394. [+ G.texensis]

Souroukis K, Cade WH, Rowell G. 1992. Factors that possibly influence variation in the calling song of field crickets: Temperature, time, and male size, age, and wing morphology. Can. J. Zool. 70: 950-955. [Gryllus texensis]

Souroukis K, Cade WH. 1993. Reproductive competition and selection on male traits at varying sex ratios in the field cricket, Gryllus pennsylvanicus. Behaviour 126: 45-62.

Souroukis K, Murray AM. 1995. Female mating behavior in the field cricket, Gryllus pennsylvanicus (Orthoptera: Gryllidae) at different operational sex ratios. J. insect Behav. 8: 269-279.

Wagner WE, Jr., Murray AM, Cade WH. 1995. Phenotypic variation in the mating preferences of female field crickets, Gryllus integer. Anim. Behav. 49: 1269-1281. [= Gryllus texensis]

Wagner WE, Jr. 1996. Convergent song preferences between female field crickets and acoustically orienting parasitoid flies. Behav. Ecol. 7: 279-285. [Gryllus lineaticeps]

Wagner WE, Jr., Hoback WW. 1999. Nutritional effects on male calling behaviour in the variable field cricket. Anim. Behav. 57: 89-95. [Gryllus lineaticeps]

Wagner WE, Jr., Reiser MG. 2000. The importance of calling song and courtship song in female mate choice in the variable field cricket. Anim. Behav. 59: 1219-1226. [Gryllus lineaticeps]

Walker TJ. 1974. Gryllus ovisopis n sp: a taciturn cricket with a life cycle suggesting allochronic speciation. Fla. Entomol. 57: 13-22.

Walker TJ. 1980. Mixed oviposition in individual females of Gryllus firmus: graded proportions of fast-developing and diapause eggs. Oecologia 47: 291-298.

Walker TJ. 1986. Monitoring the flights of field crickets, Gryllus spp., and a tachinid fly, Euphasiopteryx ochracea, in north Florida. Fla. Entomol. 69: 678-685.

Walker TJ. 1987. Wing dimorphism in Gryllus rubens, Orthoptera: Gryllidae. Ann. Entomol. Soc. Am. 80: 547-560.

Walker TJ. 1993. Phonotaxis in female Ormia ochracea (Diptera: Tachinidae), a parasitoid of field crickets. J. insect Behav. 6: 389-410.

Walker TJ. 1998. Trilling field crickets in a zone of overlap (Orthoptera: Gryllidae: Gryllus). Ann. Entomol. Soc. Am. 91: 175-184. [Gryllus rubens and G. texensis]

Walker TJ. 2000. Pulse rates in the songs of trilling field crickets (Orthoptera: Gryllidae: Gryllus). Ann. Entomol. Soc. Am. 93: 565-572. [Gryllus rubens and G. texensis]

Walker TJ. 2001. Gryllus cayensis n. sp. (Orthoptera: Gryllidae), a taciturn wood cricket extirpated from the Florida Keys: songs, ecology and hybrids. Fla. Entomol. 84: 700-705.

Walker TJ, Sivinski JM. 1986. Wing dimorphism in field crickets, Orthoptera: Gryllidae: Gryllus. Ann. Entomol. Soc. Am. 79: 84-90.

Weissman DB, Rentz DCF, Alexander RD, Loher W. 1980. Field crickets (Gryllus and Acheta) of California and Baja California Mexico (Orthoptera: Gryllidae: Gryllinae). Trans. Am. Entomol. Soc. 106: 327-356.

Weissman DB, Walker TJ, Gray DA. 2009. The field cricket Gryllus assimilis and two new sister species (Orthoptera: Gryllidae). Ann. Entomol. Soc. Am. 102(3): 367-380.

Weissman DB, Gray DA. 2019. Crickets of the genus Gryllus in the United States. Zootaxa [in manuscript to be published in May 2019]

Wiegmann DD. 1999. Search behaviour and mate choice by female field crickets, Gryllus integer. Anim. Behav. 58: 1293-1298. [=Gryllus texensis]

Willett CS, Ford MJ, Harrison RG. 1997. inferences about the origin of a field cricket hybrid zone from a mitochondrial DNA phylogeny. Heredity. Nov. 79: 484-494. [Gryllus pennsylvanicus and G. firmus]

Wineriter SA, Walker TJ. 1988. Group and individual rearing of field crickets (Orthoptera:Gryllidae). Entomol. News 99: 53-62.

Veasey JN, Kay CAR, Walker TJ, Whitcomb WH. 1976. Seasonal abundance, sex ratio, and macroptery of field crickets in northern Florida.  Ann. Entomol. Soc. Am. 69: 374-380.

Zera AJ, Rankin MA. 1989. Wing dimorphism in Gryllus rubens: Genetic basis of morph determination and fertility differences between morphs. Oecologia 80: 249-255.

Zera AJ, Mole S. 1994. The physiological costs of flight capability in wing-dimorphic crickets. Res. Popul. Ecol. (Kyoto) 36: 151-156. [Gryllus veletis and G. pennsylvanicus]

Zuk M. 1987. The effects of gregarine parasites, body size, and time of day on spermatophore production and sexual selection in field crickets. Behav. Ecol. Sociobiol. 21: 65-72. [Gryllus veletis and G. pennsylvanicus]

Zuk M. 1987. The effects of gregarine parasites on longevity, weight loss, fecundity and developmental time in the field crickets Gryllus veletis and Gryllus pennsylvanicus. Ecol. Entomol. 12: 349-354.

Zuk M. 1987. Seasonal and individual variation in gregarine parasite levels in the field crickets Gryllus veletis and Gryllus pennsylvanicus. Ecol. Entomol. 12: 341-348.

Zuk M. 1987. Age determination of adult field crickets: Methodology and field applications. Can. J. Zool. 65: 1564-1566. [Gryllus veletis and G. pennsylvanicus]

Zuk M. 1987. Variability in attractiveness of male field crickets (Orthoptera: Gryllidae) to females. Anim. Behav. 35: 1240-1248. [Gryllus veletis and G. pennsylvanicus]

Zuk M. 1988. Parasite load, body size, and age of wild-caught male field crickets (Orthoptera: Gryllidae): Effects on sexual selection. Evolution 42: 969-976. [Gryllus veletis and G. pennsylvanicus]

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