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Reproductive Biology and Diets of Goannas (Reptilia: Varanidae) from Australia Author(S): Craig D

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Reproductive Biology and Diets of Goannas (Reptilia: Varanidae) from Australia Author(S): Craig D Society for the Study of Amphibians and Reptiles Reproductive Biology and Diets of Goannas (Reptilia: Varanidae) from Australia Author(s): Craig D. James, Jonathan B. Losos and Dennis R. King Source: Journal of Herpetology, Vol. 26, No. 2 (Jun., 1992), pp. 128-136 Published by: Society for the Study of Amphibians and Reptiles Stable URL: http://www.jstor.org/stable/1564852 Accessed: 29-04-2016 03:43 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/1564852?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://about.jstor.org/terms JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Society for the Study of Amphibians and Reptiles is collaborating with JSTOR to digitize, preserve and extend access to Journal of Herpetology This content downloaded from 128.103.149.52 on Fri, 29 Apr 2016 03:43:59 UTC All use subject to http://about.jstor.org/terms Journal of Herpetology, Vol. 26, No. 2, pp. 128-136, 1992 Copyright 1992 Society for the Study of Amphibians and Reptiles Reproductive Biology and Diets of Goannas (Reptilia: Varanidae) from Australia CRAIG D. JAMES,1'4 JONATHAN B. LOSOS,2'5 AND DENNIS R. KING3 'Zoology Building A08, University of Sydney, New South Wales 2006, Australia 2Museum of Vertebrate Zoology, 1120 Life Sciences Building, University of California, Berkeley, California 94720, USA, and 3Western Australian Museum, Francis Street, Perth, Western Australia 6000, Australia ABSTRACT. - Aspects of the reproductive biology and diets of nine species of Varanus from the seasonal tropics and temperate zones of Australia were examined by dissection of 386 preserved museum specimens. We investigated whether morphological and habitat diversity in the varanids was expressed in reproductive and dietary diversity. Most species from seasonal tropical areas had enlarged testes in the dry season (June-October) and females with oviductal eggs were collected at the end of the dry season and the beginning of the wet season (September-January). However, three species seemed to be exceptions to this trend, with an extended reproductive season (V. storr), or reproductive activity in the late wet season (V. semiremex and V. kingorum). A diversity of reproductive seasons for varanids from seasonal tropical regions in Australia was also found in a review of the literature. Temperate zone species had enlarged testes in spring (September-November) and laid eggs in spring or late summer. Clutch sizes ranged from 1-7 eggs. Regression slopes of maternal SVL and clutch size seemed to be higher for the smaller-bodied species compared to the larger-bodied species, but sample sizes were too low to compare them statistically. A review of reproductive data on Australian varanids suggested that some large-bodied species (e.g., V. varius) had relatively small clutch sizes. Most species had generalized diets with lizards and orthopterans being the most important prey. Large-bodied species such as V. giganteus and V. glebopalma tended to eat larger prey items, and specialized on vertebrate prey. Varanus semiremex ate mostly aquatic prey. Diets were relatively similar for most species but appear to be influenced by body size and habitat. The family Varanidae consists of one extant ization seems to be based on meager data. Var- genus (Varanus) with about 35 species distrib- anids can be found in most major climatic uted throughout tropical Africa, southern Asia, regions in Australia (seasonal tropics, arid, semi- and Australia (Greer, 1989). Varanids are most arid, and mesic-temperate) occupying a range diverse in Australia, where 25 species range in of habitats (semi-aquatic, terrestrial and arbo- body size from the smallest living species of real). After assessing the ecological literature Varanus (V. brevicauda) at 8 g, to Australia's larg- on varanids, Shine (1986) asked, "Is this eco- est lizard, the perentie (V. giganteus), at 17 kg logical homogeneity a real phenomenon, or (Butler, 1970; Cogger, 1986; Weavers, 1988). De- simply an artifact of insufficient data?" In this spite this large range of body sizes, varanids are study we examine nine species of Australian relatively conservative in body plan (Shine, varanids in an attempt to evaluate some of the 1986; Greer, 1989), and appear also to be rela- ecological diversity of the group. We chose spe- tively similar in their ecologies: most varanids cies covering a 1000-fold range in body mass, have generalized diets, dominated by small in- and species from many different habitats and vertebrates, with a few large vertebrate prey climatic regimes, for an examination of repro- (Losos and Greene, 1988); reproductive cycles ductive biologies and diets. appear to be similar in their timing (Bustard, Only two species of Varanus, V. komodoensis 1970; King and Green, 1979; King and Rhodes, and V. olivaceus, have been studied in detail 1982; Pianka, 1986; Horn and Visser, 1989). (Auffenberg, 1981, 1988). Captive husbandry has Although the varanids appear to be superfi- provided data on the reproductive biology (e.g., cially similar in their ecologies, this general- behavior, incubation periods, clutch sizes) of a number of Australian species (Horn, 1978; Horn and Visser, 1989), but the reproductive biolo- 4 Present Address: CSIRO Division of Wildlife and gies of only six species of Australian varanids have been examined in the field (King and Ecology, P.O. Box 84, Lyneham, Australian Capitol Territory 2602, Australia. Green, 1979; King and Rhodes, 1982; Shine, 5 Present Address: Center for Population Biology, 1986). Similarly, the feeding biology of most of Storer Hall, University of California, Davis, Califor- the Australian species in the field is poorly nia 95616, USA. known. Of the species examined in this study, This content downloaded from 128.103.149.52 on Fri, 29 Apr 2016 03:43:59 UTC All use subject to http://about.jstor.org/terms REPRODUCTION AND DIETS IN VARANUS 129 no dietary data are available on four species (V. Each specimen was measured for SVL (mm), glauerti, V. kingorum, V. primordius and V. semi- blot-dried, weighed, and dissected with a mid- remex), and the diets of two other species are ventral incision. The reproductive data record- known from only single observations (V. gle- ed were: sex; state of maturity (immature or bopalma and V. pilbarensis). mature); stage of reproductive cycle (reproduc- tive or non-reproductive); level of development THE SPECIES of gonads in reproductive females (yolking We examined nine species covering almost ovarian follicles or oviductal eggs); and clutch the maximum range in body sizes of Australian sizes from females with enlarged vitellogenic varanids, and species from many different hab- follicles (>3 mm diameter), oviductal eggs or itats and climatic regions. A brief outline of the macroscopically visible corpora lutea. Stage of comparative sizes and climatic zones occupied maturity for males was judged by the appear- by the species examined is presented below. ance of the testes and epididymides, which were Maximum recorded snout-vent lengths (SVL) small and without obvious convolutions in im- for the species from largest to smallest (Greer, mature specimens. Immature females had only 1989) are V. giganteus (800 mm), V. glebopalma small follicles in the ovary (< 1.0 mm diameter) (350 mm), V. semiremex (260 mm), V. glauerti (230 and had thin, strap-like oviducts. Males were mm), V. gilleni (190 mm), V. pilbarensis (170 mm), judged to be reproductive if their testes and V. storri (130 mm), V. kingorum (120 mm), and epididymides were turgid and white, whereas V. primordius (110 mm). reproductive females were those with either The species studied are found in two major yolking ovarian follicles or oviductal eggs. Ap- climatic regions: arid and semi-arid zones (V. proximate size at sexual maturity was deter- giganteus, V. gilleni, V. pilbarensis), and seasonal mined from the overlap in the SVL of the largest tropical zones (V. glauerti, V. glebopalma, V. kin- specimen that was judged to be immature and gorum, V. primordius, V. semiremex and V. storri). the smallest reproductive specimen. The seasonal tropics in northern Australia (north Stomach contents were removed and inver- of 20?S latitude) have high year-round diurnal tebrate prey items were identified to order, temperatures (>30 C) with strongly seasonal whereas vertebrate prey items were identified rainfall (wet-dry topics). Monsoonal rainfall be- to species if possible. Substantially intact prey gins in December and lasts until April. From items were blot-dried and weighed. The mass May until November little or no rainfall is re- ratio (MR = prey mass/predator mass) was cal- ceived and the region becomes increasingly dry culated for each intact prey item (Greene, 1986; (see James and Shine, 1985, for more details of Losos and Greene, 1988). Counts of fragmentary the climate). In the arid and semi-arid regions body parts were used to estimate the minimum of central Australia, rainfall is low (around 250 number of individual prey. Fragmented prey mm annual average) and unpredictable, and items were assigned the mean MR for that prey there is a marked difference between daily sum- taxon in that species of Varanus. The Importance mer and winter temperatures. Index (II, hereafter) for each prey taxa for a Most of the species from the seasonal tropics species is the sum of MRs for that prey taxa occur in rocky habitats (V. glauerti, V. glebopalma, divided by all MRs for that species (Greene, V.
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