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Variation in testis weight of the Tibetan toad Scutiger boulengeri along a narrow altitudinal gradient

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Life-history theory predicts that organisms inhabiting harsh environments such as high altitudes should invest less in reproduction and more in survival. Testis size is associated with the intensity of male-male competition for mating and thus may be treated as an indicator of male reproductive investment. Hence, it may be expected that organisms will reduce their testis size with increasingly harsh environments. Here we test this prediction in a toad species, Scutiger boulengeri, endemic to the Tibetan plateau using data from three populations located at altitudes of 4078, 4276, and 4387 m. Consistent with the prediction, male toads exhibited smaller testes at higher altitudes, despite the relatively narrow altitudinal span. It is likely that cold climates and strong seasonality constrain the ability of high-altitude male toads to allocate more energy into reproduction, thereby leading to small testis size. In addition, the left testis was significantly heavier than the right one and the degree of size asymmetry was unrelated to either altitude or body condition.

Affiliations: 1: 1Department of Ecology, College of Life Sciences, Henan Normal University, Xinxiang 453007, China ; 2: 2Department of Ecology, College of Life Sciences, Wuhan University, Wuhan 430072, China

*Corresponding author; e-mail:

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1. Berven K.A. (1982) "The genetic basis of altitudinal variation in the wood frog Rana sylvatica. I. An experimental analysis of life history traits". Evolution, Vol 36, 962-983.
2. Briskie J.V., Montgomerie R. (2007) "Testis size, sperm size and sperm competition". In: Jamieson B.G.M. (Ed.) Reproductive Biology and Phylogeny of Birds. Part A: Phylogeny, Morphology, Hormones, Fertilization, pp.  513-551. Science Publishers, Enfield, NH, USA.
3. Brown J.L. (1964) "The evolution of diversity in avian territorial systems". Wilson Bull., Vol 76, 160-169.
4. Byrne P., Roberts J.D., Simmons L.W. (2002) "Sperm competition selects for increased testis mass in Australian frogs". J. Evol. Biol., Vol 15, 347-355. [Crossref]
5. Calhim S., Birkhead T.R. (2007) "Testes size in birds: quality versus quantity-assumptions, errors, and estimates". Behav. Ecol., Vol 18, 271-275. [Crossref]
6. Chen W., Tang Z.H., Fan Y.G., Wang Y., Pike D.A. (2013) "Maternal investment increases with altitude in a frog on the Tibetan Plateau". J. Evol. Biol., Vol 26, 2710-2715. [Crossref]
7. Chen W., Pike D.A., He D.J., Wang Y., Ren L.N., Wang X.Y., Fan X.G., Lu X. (2014) "Altitude decreases testis weight of a frog (Rana kukunoris) on the Tibetan Plateau". Herpetol. J., Vol 24, 183-188.
8. Clutton-Brock T.H., Hodge S.J., Flower T.P. (2008) "Group size and the suppression of subordinate reproduction in Kalahari meerkats". Anim. Behav., Vol 76, 689-700. [Crossref]
9. Cummins C.P. (1986) "Temporal and spatial variation in egg size and fecundity in Rana temporaria". J. Anim. Ecol., Vol 55, 303-316. [Crossref]
10. Czarnołe’ski M., Kozłowski J. (1998) "Do Bertalanffy’s growth curves result from optimal resource allocation?" Ecol. Lett., Vol 1, 5-7. [Crossref]
11. Duellman W.E., Trueb L. (1986) Biology of Amphibians. McGraw-Hill Book Company, Toronto, ON, Canada.
12. Dziminski M.A., Roberts J.D., Beveridge M., Simmons L.W. (2009) "Sperm competitiveness in frogs: slow and steady wins the race". Proc. R. Soc. B, Vol 276, 3955-3961. [Crossref]
13. Emlen S.T., Oring L.W. (1977) "Ecology, sexual selection, and the evolution of mating systems". Science, Vol 197, 215-223. [Crossref]
14. Fei L., Hu S.Q., Ye C.Y., Tian W.S., Jiang J.P., Zhong S.X., Wang Y.S. (2009) Fauna Sinica, Amphibia Vol. 3, Anura Ranidae. Science Press, Beijing, China.
15. Gibbons M.M., McCarthy T.K. (1986) "The reproductive output of frogs Rana temporaria (L.) with particular reference to body size and age". J. Zool., Vol 209, 579-593. [Crossref]
16. Gollman B., Gollman G. (1996) "Geographic variation of larval traits in the Australian frog Geocrinia victoriana". Herpetologica, Vol 52, 181-187.
17. Grant J.W.A., Gaboury C.L., Levitt H.L. (2000) "Competitor-to-resource ratio, a general formulation of operational sex ratio, as a predictor of competitive aggression in Japanese medaka (Pisces: Oryziidae)". Behav. Ecol., Vol 11, 670-675. [Crossref]
18. Halliday T.R., Verrell P.A. (1988) "Body size and age in amphibians and reptiles". J. Herpetol., Vol 22, 253-265. [Crossref]
19. Harcourt A.H., Harvey P.H., Larson S.G., Short R.V. (1981) "Testis weight, body weight, and breeding system in primates". Nature, Vol 293, 55-57. [Crossref]
20. Hettyey A., Laurila A., Herczeg G., Jönsson K.I., Kovács T., Merilä J. (2005) "Does testis weight decline towards the Subarctic? A case study on the common frog, Rana temporaria". Naturwissenschaften, Vol 92, 188-192. [Crossref]
21. Howard J.H., Wallace R.L. (1985) "Life history characteristics of populations of the long-toed salamander (Ambystoma macrodactylum) from different altitudes". Am. Midl. Nat., Vol 113, 361-372. [Crossref]
22. Jin L., Mi Z.P., Liao W.B. (2016a) "Altitudinal variation in male reproductive investment in a polyandrous frog species (Hyla gongshanensis jingdongensis)". Anim. Biol., Vol 66, 289-303. [Crossref]
23. Jin L., Yang S.N., Liao W.B., Lüpold S. (2016b) "Altitude underlies variation in the mating system, somatic condition and investment in reproductive traits in male Asian grass frogs (Fejervarya limnocharis)". Behav. Ecol. Sociobiol., Vol 70, 1197-1208. [Crossref]
24. Kappeler P.M. (1997) "Intrasexual selection and testis size in strepsirhine primates". Behav. Ecol., Vol 8, 10-19. [Crossref]
25. Koskela P., Pasanen S. (1975) "The reproductive biology of the female common frog, Rana temporaria L., in northern Finland". Aquilo Ser. Zoologica., Vol 16, 1-12.
26. Lake P.E. (1981) "Male genital organs". In: King A.S., McLelland J. (Eds) Form and Function in Birds, pp.  1-61. Academic Press, London, UK.
27. Liao W.B., Lu X. (2010) "Age and growth of a subtropical high-elevation torrent frog, Amolops mantzorum, in western China". J. Herptol., Vol 44, 172-176. [Crossref]
28. Liao W.B., Lu X. (2012) "Adult body size=f(initial size+growth rate×age): explaining the proximate cause of Bergman’s cline in a toad along altitudinal gradients". Evol. Ecol., Vol 26, 579-590. [Crossref]
29. Liao W.B., Lu X., Shen Y.W., Hu J.C. (2011) "Age structure and body size of two populations of the rice frog Rana limnocharis from different altitudes". Ital. J. Zool., Vol 78, 215-221. [Crossref]
30. Liao W.B., Lou S.L., Zeng Y., Kotrschal A. (2016) "Large brains, small guts: the expensive tissue hypothesis supported in anurans". Am. Nat., Vol 188, 693-700. [Crossref]
31. Licht L.E. (1975) "Comparative life history features of the western spotted frog, Rana pretiosa, from low- and high-elevation populations". Can. J. Zool., Vol 53, 1254-1258. [Crossref]
32. Liu Y.H., Liao W.B., Zhou C.Q., Mi Z.P., Mao M. (2011) "Asymmetry of testes in Guenther’s frog, Hylarana guentheri (Anuar: Ranidae)". Asian Herpetol. Res., Vol 2, 234-239.
33. Lu X., Li B., Liang J.J. (2006) "Comparative demography of a temperate anuran, Rana chensinensis, along a relatively fine elevational gradient". Can. J. Zool., Vol 84, 1789-1795. [Crossref]
34. Ma X.Y., Lu X. (2009) "Sexual size dimorphism in relation to age and growth based on skeletochronological analysis in a Tibetan frog". Amphibia-Reptilia, Vol 30, 351-359. [Crossref]
35. Ma X.Y., Lu X., Merilä J. (2009a) "Altitudinal decline of body size in a Tibetan frog". J. Zool., Vol 279, 364-371. [Crossref]
36. Ma X.Y., Tong L.N., Lu X. (2009b) "Variation of body size, age structure and growth of a temperate frog, Rana chensinensis, over an elevational gradient in northern China". Amphibia-Reptilia, Vol 30, 111-117. [Crossref]
37. Mai C.L., Liu Y.H., Jin L., Mi Z.P., Liao W.B. (2017) "Altitudinal variation in somatic condition and investment in reproductive traits in male Yunnan pond frog (Pelophylax pleuraden)". Zool. Anz., Vol 266, 189-195. [Crossref]
38. Merilä J., Sheldon B.C. (1999) "Testis size variation in the greenfinch Carduelis chloris: relevance for some recent models of sexual selection". Behav. Ecol. Sociobiol., Vol 45, 115-123. [Crossref]
39. Mi Z.P., Liao W.B., Jin L., Lou S.L., Cheng J., Wu H. (2012) "Testes asymmetry and sperm length in Rhacophorus omeimontis". Zool. Sci., Vol 29, 368-372. [Crossref]
40. Møller A.P. (1994) "Directional selection and directional asymmetry: testes size and secondary sexual characters in birds". Proc. R. Soc. B, Vol 258, 147-151. [Crossref]
41. Morrison C., Hero J.M. (2003) "Geographic variation in life history characteristics of amphibians: a review". J. Anim. Ecol., Vol 72, 270-279. [Crossref]
42. Parker G.A. (1970) "Sperm competition and its evolutionary consequences in the insects". Biol. Rev., Vol 45, 525-567. [Crossref]
43. Pitcher T.E., Stutchbury B.J.M. (1998) "Latitudinal variation in testis size in six species of North American songbirds". Can. J. Zool., Vol 76, 618-622. [Crossref]
44. Pitcher T.E., Dunn P.O., Whittingham L.A. (2005) "Sperm competition and the evolution of testes size in birds". J. Evol. Biol., Vol 18, 557-567. [Crossref]
45. Rising J.D. (1996) "Relationship between testis size and mating systems in American sparrows (Emberizinae)". Auk, Vol 113, 224-228. [Crossref]
46. Roff D.A. (2002) Life History Evolution. Sinauer Associates Inc., Sunderland, MA, USA.
47. Ryser J. (1996) "Comparative life histories of a low- and a high-elevation population of the common frog Rana temporaria". Amphibia-Reptilia, Vol 17, 183-195. [Crossref]
48. Schulte-Hostedde A.I., Millar J.S. (2004) "Intraspecific variation of testis size and sperm length in the yellowpine chipmunk (Tamias amoenus): implications for sperm competition and reproductive success". Behav. Ecol. Sociobiol., Vol 55, 272-277. [Crossref]
49. Sehulte-Hostedde A.I., Millar J.S., Hiekling G.J. (2005) "Condition dependence of testis size in small mammals". Evol. Ecol. Res., Vol 7, 143-149.
50. Simmons L.W., Kotiaho J.S. (2002) "Evolution of ejaculates: patterns of phenotypic and genotypic variation and condition dependence in sperm competition traits". Evolution, Vol 56, 1622-1631. [Crossref]
51. Song Z.M., Huang D.M., Chang C. (1990) "On the development and population age structure of Scutiger boulengeri tadpoles". Acta Zool. Sin., Vol 36, 187-192.
52. Tang T., Luo Y., Huang C.H., Liao W.B., Huang W.C. (2018) "Variation in somatic condition and testes mass in Feirana quadranus along an altitudinal gradient". Anim. Biol. DOI:10.1163/15707563-17000142.
53. Tilley S.G. (1980) "Life histories and comparative demography of two salamander populations". Copeia, Vol 1980, 806-821. [Crossref]
54. Zamudio K.R., Bell R.C., Nali R.C., Haddad C.F.B., Prado C.P.A. (2016) "Polyandry, predation, and the evolution of frog reproductive modes". Am. Nat., Vol 188, S41-S61. [Crossref]
55. Zeng Y., Lou S.L., Liao W.B., Jehle R. (2014) "Evolution of sperm morphology in anurans: insights into the roles of mating system and spawning locations". BMC Evol. Biol., Vol 14, 104. DOI:10.1186/1471-2148-14-104. [Crossref]
56. Zhang L.X., Lu X. (2012) "Amphibians live longer at higher altitudes but not at higher latitudes". Biol. J. Linn. Soc., Vol 106, 623-632. [Crossref]
57. Zhang L.X., Ma X.Y., Jiang J.P., Lu X. (2012) "Stronger condition dependence in female size explains altitudinal variation in sexual size dimorphism of a Tibetan frog". Biol. J. Linn. Soc., Vol 107, 558-565. [Crossref]
58. Zhou C.Q., Mao M., Liao W.B., Mi Z.P., Liu Y.H. (2011) "Testis asymmetry in the dark-spotted frog Rana nigromaculata". Herpetol. J., Vol 21, 181-185.

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