Allometry of clutch and neonate sizes in monitor lizards (Varanidae: Varanus)

Copeia

Volumn , Issue 2, 2001, Pages 443-458

  Thompson, Graham G ^a   Pianka, Eric R ^b  

^a EDITH COWAN UNIVERSITY   (Australia)

^b UNIVERSITY OF TEXAS AT AUSTIN   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ALLOMETRY; BODY SIZE; CLUTCH SIZE; LIZARD; REPRODUCTIVE BIOLOGY;

VARANUS;

EID: 0035339678     PISSN: 00458511     EISSN: None     Source Type: Journal    
DOI: 10.1643/0045-8511(2001)001[0443:AOCANS]2.0.CO;2     Document Type: Article

References (122)

1. ANDREWS, H. V. 1995. Sexual maturation in Varanus salvator (Laurenti, 1768), with notes on growth and reproductive effort. Herpetol. J. 5:189-194.
2. _, AND M. GAULKE. 1990. Observations on the reproductive biology and growth of the water monitor (Varanus salvator) at the Madras Crocodile Bank. Hamadryad 15:1-5.
3. ANDREWS, R. M., AND A. S. RAND. 1974. Reproductive effort in anoline lizards. Ecology 55:1317-1327.
4. ANONYMOUS. 1995. Gazella. News. Zoo Praha. Vol. 22, Prague, Czech Republic.
5. AUFFENBERG, W. 1981. The behavioral ecology of the Komodo monitor. Univ. Presses of Florida, Gainesville.
6. _. 1988. Gray's monitor lizard. Univ. Presses of Florida, Gainesville.
7. _. 1994. The Bengal monitor. Univ. Presses of Florida, Gainesville.
8. _, H. RAHMAN, F. IFFAT, AND Z. PERVEEN. 1989. A study of Varanus flavescens (Harwicke & Gray) (Sauria: Varanidae). J. Bombay Nat. Hist. Soc. 86: 286-307.
9. BARBOUR, T., AND A. LOVERIDGE. 1928. A comparative study of the herpetological fauna of the Uluguru and Usambara mountains, Tanganyika territory, with descriptions of new species. Mem. Mus. Comp. Zool., Cambridge, MA, 50:87-265.
10. BARNETT, B. 1979. Incubation of sand goanna (Varanus gouldii) eggs. Herpetofauna 11:21-22.
11. BARTLETT, R. D. 1982. Initial observations on the captive reproduction of Varanus storri, Mertens. Ibid. 13(2):6-7.
12. BAVERSTOCK, P. R., D. KING, M. KING, J. BIRRELL, AND M. KRIEG. 1993. The evolution of species of the Varanidae: microcomplement fixation analysis of serum albumins. Aust. J. Zool. 41:621-638.
13. BAYLESS, M. 1992. Notes on the reproductive behavior of the Nile monitor lizard Varanus niloticus Linnaeus (1766). Varanews 2:5-6.
14. _. 1992. Reproductive notes on the black roughneck monitor lizard (Varanus rudicollis Gray, 1845). Ibid. 3:3.
15. _. 1994. Zur fortpfanzungsbiologie des steppenwarans (Varanus exanthematicus). Salamandra 30:109-118.
16. _, AND R. HUFFAKER. 1992. Observations of egg deposition and hatching of the savannah monitor (Varanus exanthematicus Bocs, 1792) in captivity. Varanews 3:5-6.
17. _, AND T. REYNOLDS. 1992. Breeding of the savannah monitor lizard in captivity (Varanus exanthematicus Bocs, 1972). Herpetology 22:12-14.
18. _, R. HUFFAKER, AND O. MAERCPLS. 1994. Notes on the egg deposition and incubation of the argus monitor (Varanus gouldii horni, Gray 1838) in captivity. Varanews 4:5.
19. BIRCHARD, G.F., T. WALSH, R. ROSSCOE, AND C. L. REIBER. 1995. Oxygen uptake by Komodo dragon (Varanus komodoensis) eggs: the energetics of prolonged development in a reptile. Physiol. Zool. 68: 622-633.
20. BLUEWEISS, L., H. FOX, V. KUDZMA, D. NAKASHIMA, R. PETERS, AND S. SAMS. 1978. Relationships between body size and some life history parameters. Oecologia 37:257-272.
21. BOSCH, H. 1999. Successful breeding of the Emerald monitor (Varanus p. prasinus) in the Lobbecke Museum + Aquazoo, Dusseldorf (Germany). Mertensiella 11:225-226.
22. BOYLAN, T. 1995. Field observations, captive breeding and growth rates of the Lace monitor, Varanus varius. Herpetofauna 25:10-14.
23. BOYLE, D. M., AND W. F. LAMOREAUX. 1983(1984). Captive reproduction in the pygmy mulga monitor Varanus gilleni at Dallas Zoo. Proc. Rept. Symp. Capt. Prog. Husbandry 7:59-63.
24. BRANCH, W. R. 1991. The regenia registers of "Gogga" Brown (1869-1909) "Memoranda on a species of Monitor or Varan." Mertensiella 2:57-110.
25. _, AND H.-G. HORN. 1987. Uber die nachzucht des australischen risenwarans Varanus giganteus (Gray, 1845). Salamandra 23:90-96.
26. BREDL, J., AND T. D. SCHWANER. 1983. First record of captive propagation of the lace monitor, Varanus varius (Sauria: Varanidae). Herpetofauna 15:20-21.
27. BROCKELMAN, W. Y. 1975. Competition, the fitness of offspring, and optimal clutch size. Am. Nat. 109: 677-699.
28. CALDER, W. A. 1984. Size, function and life history. Harvard Univ. Press, Cambridge, MA.
29. CARLZEN, G. 1982. Breeding green tree monitors. Herpetology 12:4-6.
30. CARTER, D. B. 1999. Reproductive cycle of the lace monitor Varanus varius. Mertensiella, 11:129-135.
31. CHARNOV, E. L. 1993. Life-history Invariants. Oxford Univ. Press, London.
32. CHARNOV, E. L., AND J. F. DOWNHER. 1995. A trade-off-invariant life-history rule for optimal offspring size. Nature 376:418-419.
33. CHIPPINDALE, P. 1991. Captive breeding of the Timor monitor (Varanus timorensis similis). Herpetol. Rev. 22:52-53.
34. CHRISTIAN, A., AND T. GARLAND. 1996. Scaling of limb proportions in monitor lizards (Squamata: Varanidae). J. Herpetol. 30:219-230.
35. CHRISTIAN, T. 1981. Varanus tristis - a variable monitor. Herpetofauna 12:7-12.
36. CONGDON, J. D., AND J. W. GIBBONS. 1985. Egg components and reproductive characteristics of turtles: relationships to body size. Herpetologica 41:194-205.
37. COWLES, R. B. 1930. The life history of Varanus niloticus (Linnaeus) as observed in Natal South Africa. J. Entomol. Zool. 22:3-31.
38. DE LISLE, H. F. 1996. The natural history of monitor lizards. Krieger, Malabar, FL.
39. DOLES, M., AND W. CARD. 1995. Delayed fertilization in the monitor lizard Varanus gouldii. Herpetol. Rev. 26:196.
40. DWYER, Q., AND M. K. BAYLESS. 1996. Notes on the husbandry and reproductive behavior of the argus monitor (Varanus gouldii horni). Adv. Herpetoculture, p 81-85.
41. EHMANN, H., G. SWAN, G. SWAN, AND B. SMITH. 1991. Nesting, egg incubation and hatching by the heath monitor Varanus rosenbergi in a termite mound. Herpetofauna 21:17-24.
42. EIDENMÜLLER, B. 1986. Beobachtungen bei der pflege und nachzucht von Varanus (Odatria) t. timorensis (Gray, 1831). Salamandra 22:157-161.
43. _. 1989. Beobachtungen bei der haltung und nachzucht von Varanus (Odatria) tristis orientalis Fry, 1913. Ibid. 25:265-271.
44. _. 1985 Captive breeding of Varanus storri. Ibid. 21:55-61.
45. _. 1994. Bemerkungen zur haltung und zucht von Varanus acanthurus Boulenger, 1885, V. storri Mertens, 1966 und V. gilleni Lucus and Frost, 1985. Herpetofauna 16:6-12.
46. _. 1998. Bemerkungen zur haltung und nachzucht von Varanus p. prasinus (Schlegel, 1939) und Varanus p. beccarii (Doria, 1874). Ibid. 20:8-13
47. _, AND H.-G. HORN. 1985. Eigene nachzuchten und der gegenwartige stand der nachzucht von Varanus (Odatria) storri Mertens, 1996. Salamandra 21: 55-61.
48. _, AND R. WICKER. 1991. Einige beobachtungen bei der pflege und nachzucht von Varanus (Odatria) timorensis similis Mertens, 1958. Ibid. 27:187-193.
49. _, AND _. 1995. The successful breeding of Mertens' monitor lizard, Varanus mertensi, Glauerti 1951. Herpetofauna 25:4-7.
50. ELLIS, C. 1998. The land of the Komodo dragon. Times Ed. Singapore.
51. FELSENSTEIN, J. 1985. Phytogenies and the comparative method. Am. Nat. 125:1-15.
52. FITCH, H. S. 1970. Reproductive cycles in lizards and snakes. Univ. Kans. Mus. Nat. Hist. Misc. Publ. 52: 1-247.
53. _. 1985. Variation in clutch and litter size in New World reptiles. Ibid. 76:1-76.
54. FORD, N., AND R. A. SEIGEL. 1989. Relationships among body size, clutch size, and egg size in three species of oviparous snakes. Herpetologica 45:75-83.
55. FILLER, S., P. BAVERSTOCK, AND D. KING. 1998. Biogeographic origins of goannas (Varanidae); a molecular perspective. Mol. Phyl. Evol. 9:294-307.
56. GARLAND, T., A. W. DICKERMAN, C. M. JANIS, AND J. A. JONES. 1993. Phylogenetic analysis of covariance by computer simulation. Syst. Biol. 42:265-292.
57. GOW, G. F. 1982. Notes on the reproductive biology of the pygmy mulga goanna Varanus gilleni Lucas & Frost, 1895. Northern Territory Nat. 1:19-20.
58. GREEN, B., M. MCKELVEY, AND P. RISMILLER. 1999. The behaviour and energetics of hatchling Varanus rosenbergi. Mertensiella 11:105-112.
59. GREENE, H. W. 1986. Diet and Arboreality in the Emerald Monitor, Varanus prasinus, with comments on the study of adaptation. Fieldiana 31:1-12.
60. GREER, A. E. 1989. Biology and evolution of Australian lizards. Surrey Beatty, Sydney, New South Wales, Australia.
61. GUPTA, B. K. 1996. A note on reproduction in the desert monitor (Varanus griseus). Hamadryad 21:43.
62. HAIRSTON, C., AND P. BURCHFIELD. 1990. A sustained captive breeding program for a large varanid lizard Varanus salvator at the Gladys Porter Zoo, p. 22-28. In: AAZPA Regional Conference Proceedings American Association for Zoological Parks and Aquariums, Wheeling, WV.
63. _, AND _. 1992. The reproduction and husbandry of the water monitor Varanus salvator at the Gladys Porter Zoo, Brownsville. Int. Zoo Yb. 31: 124-130.
64. HARVEY P. H., AND M. D. PAGEL. 1991. The comparative method in evolutionary biology. Oxford Univ. Press, Oxford.
65. HERMANN, H.-W. 1999. Husbandry and captive breeding of the water monitor, Varanus salvator (Reptilia: Sauria: Varanidae) at the Cologne Aquarium (Cologne Zoo). Mertensiella 11:95-103.
66. HORN, H. G. 1978. Nachzucht von Varanus gilleni (Reptilia: Sauria: Varanidae). Salamandra 14:29-32.
67. _, AND G. J. VISSER. 1989. Review of reproduction of monitor lizards Varanus spp. in captivity. Int. Zoo Yb. 28:140-150.
68. _, AND _. 1991. Basic data on the biology of monitors. Mertensiella 2:176-187.
69. _, _. 1997. Review of reproduction of monitor lizards Varanus spp in captivity II. Int. Zoo Yb. 35:227-246.
70. HUSBAND, G. A. 1980. Notes on a nest and hatchlings of Varanus acanthurus. Herpetofauna (Sydney) 11: 29-30.
71. _. 1989. Captive breeding of the pigmy mulga monitor at the Australian Reptile Park. Aust. Herpetol. News 7:10.
72. IRWIN, B. 1986. Captive breeding of two species of monitor. Thylacinus 11:4-5.
73. IRWIN, S. 1996a. Capture, field observations and husbandry of the rare canopy goanna. Ibid. 21:12-19.
74. _. 1996b. Courtship, mating and egg deposition by the captive perentie Varanus giganteus at Queensland Reptile and Fauna Park. Ibid. 21:8-11.
75. JAMES, C. 1996. Ecology of the pygmy goanna (Varanus brevicauda) in spinifex grasslands of central Australia. Aust. J. Zool. 44:177-192.
76. _, AND W. G. WHITFORD. 1994. An experimental study of phenotypic plasticity in the clutch size of a lizard. Oikos 70:49-56,
77. _, J. B. LOSOS, AND D. R. KING. 1992. Reproductive biology and diets of goannas (Reptilia: Varanidae) from Australia. J. Herpetol. 26:128-136.
78. KING, D., AND B. GREEN. 1979. Notes on diet and reproduction of the sand goanna Varanus gouldii rosenbergi. Copeia 1979:64-70.
79. _, AND _. 1999. Goanna: the biology of Varanid lizards. New South Wales Univ. Press, Sydney, New South Wales, Australia.
80. _, AND L. RHODES. 1982. Sex ratio and breeding season of Varanus acanthurus. Copeia 1982:784-787.
81. _, B. GREEN, AND H. BUTLER. 1989. Activity patterns, temperature regulation and diet of Varanus giganteus on Barrow Island, Western Australia. Aust. Wildl. Res. 16:41-47.
82. KLUGE, A. G. 1987. Cladistic relationships in the gekkonoidae (Squamata, Sauria). Misc. publ. Mus. Zool. Univ. Mich. 174:1-54.
83. KREBS, U. 1999. Experimental variation of breeding season and incubation time in the spiny-tailed monitor (Varanus acanthurus). Mertensiella 11:227-237.
84. MADSEN, T., AND R. SHINE. 1996. Determinants of reproductive output in female water pythons (Liasis fuscus: Pythonidea). Herpetologica 52:146-159.
85. MARKWELL, K. 1983. The artificial incubation of Lace monitor (Varanus varius) eggs. Herpetogauna 15: 16-17.
86. MITCHELL, L. A. 1989. Reproduction of Gould's monitor (Varanus gouldii) at the Dallas Zoo. Bull. Chiago Herpetol. Soc. 25:8-9.
87. NABORS, P. 1997. Notes on breeding the argus monitor, Varanus panoptes, in captivity. Dragon News 1: 3-4.
88. PENGILLEY, R. 1981. Notes on the biology of Varanus spenceri and V. gouldii, Barkly Tablelands, Northern Territory. Aust. J. Herpetol. 1:23-26.
89. PERRY, G., R. HABANI, AND H. MENDELSSOHN. 1993. The first captive reproduction of the desert monitor Varanus griseus griseus at the Research Zoo of Tel Aviv University. Int. Zoo Yb. 32:188-190.
90. PETERS, U. W. 1971. The first hatching of Varanus spenceri in captivity. Bull. Zoo Man. 3:17-18.
91. PHILLIPS, J. A., AND R. P. MILLAR. 1998. Reproductive biology of the white-throated savanna monitor, Varanus albigularis. J. Herpetol. 32:366-377.
92. _, AND G. C. PACKARD. 1994. Influence of temperature and moisture on eggs and embryos of the white-throated savanna monitor Varanus albigularis: implications for conservation. Biol. Conserv. 69: 131-136.
93. PIANKA, E. R. 1970. Comparative autecology of the lizard Cnemidophorus tigris in different parts of its geographic range. Ecology 51:703-720.
94. _. 1976. Natural selection of optimal reproductive tactics. Am. Zool. 16:775-784.
95. _. 1994. Comparative ecology of Varanus in the Great Victoria Desert. Aust. J. Ecol. 19:395-408.
96. _. 1995. Evolution of body size: Varanid lizards as a model system. Am. Nat. 146:398-414.
97. QUALLS, C. P., AND R. SHINE. 1995. Maternal body-volume as a constraint on reproductive output in lizards: evidence from the evolution of viviparity. Oecologia 103:73-78.
98. RILEY, J., A.F. STIMSON, AND J. M. WINCH. 1985. A review of squamata ovipositing in ant and termite nests. Herpetol. Rev. 16:38-43.
99. SCHMIDA, G. 1974. Der kurzschwanzwaran (Varanus brevicauda). Aquar.-u. Terr. Z. 27:390-394.
100. SEIGEL, R. A., AND H. S. FITCH. 1984. Ecological patterns of relative clutch mass in snakes. Oecologia 61:293-301.
101. _, AND N. B. FORD. 1987. Reproductive ecology, p. 210-252. In: Snakes: ecology and evolutionary biology. R. A. Seigel, J. T. Collins, and S. S. Novak. (eds.). Macmillan, New York.
102. SHINE, R. 1978. Growth rates and sexual maturation in six species of Australian elapid snakes. Herpetologica 34:73-79.
103. _. 1986. Food habits, habitats and reproductive biology of four sympatric species of varanid lizards in tropical Australia. Ibid. 42:346-360.
104. _. 1992. Relative clutch mass and body shape in lizards and snakes: is reproductive investment constrained or optimized? Evolution 46:828-833.
105. _, AND A. E. GREER. 1991. Why are clutch sizes more variable in some species than others? Ibid. 45: 1696-1706.
106. _, AND L. SCHWARZKOPF. 1992. The evolution of reproductive investment in lizards and snakes. Ibid. 46:62-75.
107. _, AMBARIYANTO, P. S. HARLOW, AND MUMPUNI 1998. Ecological traits of commercially harvested water monitors, Varanus salvator, in northern Sumatra. Wildl. Res. 25:437-447.
108. SMITH, H., G. SINELNICK, J. D. FAWCETT, AND R. E. JONES. 1973. A survey of the chronology of ovulation in anoline lizard genera. Trans. Kans. Acad. Sci. 75:107-120.
109. SMITH, L. A. 1988. Notes on clutch of monitor (Varanus caudolineatus) eggs. West. Aust. Nat. 17:96.
110. STEARNS, S. C. 1992. The evolution of life histories. Oxford Univ. Press, New York.
111. STORR, G. M. 1980. The monitor lizards (genus Varanus Merrem 1820) of Western Australia. Rec. West. Aust. Mus. 8:237-293.
112. THISSEN, R. 1992. Breeding the spiny-tail monitor (Varanus acanthurus Boulenger). Vivarium 3:32-34.
113. THOMPSON, G. G. 1996. Breeding Varanus brevicauda. West. Aust. Nat. 21:119-121.
114. _, AND E. R. PIANKA. 1999. Reproductive ecology of Varanus tristis (Squamata: Varanidae). J. R. Soc. W.A. 82:27-31.
115. _, AND P. C. WITHERS. 1997. Comparative morphology of Western Australian varanid lizards (Squamata: Varanidae). J. Morphol. 233:127-152.
116. VAN DAMME, R., D. BAUWENS, F. BRANA, AND R. F. VERHEYEN. 1992. Incubation temperature differentially affects hatching time, egg survival, and hatchling performance in the lizard Podarcis muralis. Herpetologica 48:220-228.
117. VAN DUINEN, J. J. 1983. Varanenkweek in het noorderdierenpark te emmen. Lacerta 42:12-14.
118. VITT, L. J., AND J. D. CONGDON. 1978. Body shape, reproductive effort, and relative clutch mass in lizards: resolution of a paradox. Am. Nat. 112:595-608.
119. WALSH, T. 1993. Dragon tales, the history, husbandry, and breeding of Komodo monitors at the National Zoological Park. Vivarium 4:23-26.
120. WERNER, Y. L. 1989. Egg size and egg shape in Near-Eastern gekkonid lizards. Isr. J. Zool. 35:199-213.
121. WICKER, R., M. GAULKE, H.-G. HORN. 1999. Contributions to the biology, keeping and breeding of the Mindanao water monitor (Varanus s. cumingi). Mertensiella 11:213-148.
122. ZAR, J. H. 1984. Biostatistical analysis. Prentice Hall, London.