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Patterns of Predation and Antipredator Behavior in the Australian Water Dragon, Physignathus Lesueurii

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Patterns of Predation and Antipredator Behavior in the Australian Water Dragon, Physignathus Lesueurii Herpetological Conservation and Biology 9(1):48−56. Submitted: 5 November 2012; Accepted: 26 August 2013; Published: 13 July 2014. PATTERNS OF PREDATION AND ANTIPREDATOR BEHAVIOR IN THE AUSTRALIAN WATER DRAGON, PHYSIGNATHUS LESUEURII 1 2 3 4 5 J. SEAN DOODY , PETER HARLOW , DONNA DOUGLASS , JASON D. THIEM , BEN BROADHURST , 6 7 7 8 DANE TREMBATH , JERRY OLSEN , ESTEBAN FUENTES , AND TONY ROSE 1Department of Ecology and Evolutionary Biology, 569 Dabney Hall, University of Tennessee, Knoxville, Tennessee 37996- 1610, USA, e-mail: [email protected] 2Taronga Zoo, PO Box 20, Bradleys Head Road, Mosman, New South Wales 2088, Australia 3Australian National Botanical Gardens, GPO Box 1777, Canberra, Australian Capital Territory 2601, Australia 4Fish Ecology and Conservation Physiology Laboratory, Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6, Canada 5Parks, Conservation, and Lands, ACT Department of Territory and Municipal Services, GPO BOX 158, Canberra, Australian Capital Territory 2601, Australia 6Museum and Art Gallery of the Northern Territory, GPO Box 4646, Darwin Northern Territory 0801, Australia 7Institute for Applied Ecology, University of Canberra, Australian Capital Territory 2601, Australia 8The Australian Museum, 6 College Street, Sydney, New South Wales 2010, Australia Abstract.—Although lizards are more diverse in arid environments, many lizard taxa have independently invaded aquatic habitats. Adaptations in aquatic lizards are often straightforward (e.g., dorsolaterally compressed tails facilitate swimming), but what made these species invade aquatic habitats in the first place? Although this question is not directly testable, one possible reason is to reduce predation risk. Here we examine the fit of a species’ antipredator behaviors to the predation risk associated with its known and suspected predators. We reviewed the literature for records of predation on the Australian Water Dragon, Physignathus lesueurii, a semi-aquatic lizard that uses aquatic escape and aquatic sleeping, but does not forage underwater. We then examined these two behaviors in relation to the patterns of predation revealed by our review. Our review identified 25 species of predators, most of which were aerial (mainly raptors) or terrestrial/arboreal (snakes, lizards, and mammals). Aquatic predators were rare. Our review supports the hypothesis that Water Dragons invaded and persist in aquatic habitats to decrease predation risk from terrestrial and aerial predators, although we cannot rule out the exploitation of an abundance of available (riparian) food as the primary reason for invading aquatic habitats. Future studies should test how well Water Dragons can thermoregulate by sleeping underwater (vs. in air), and could test the antipredator responses of Water Dragons to mock predators. Key Words.—Agamidae; aquatic escape; aquatic lizards; aquatic sleeping; predation risk INTRODUCTION revealed 73 lizard species from 11 families that regularly use aquatic habitats, noting the multiple, independent Predator escape is an essential, adaptive behavior in evolution of aquatic or semi-aquatic species. Use of the animals, and researchers have devoted considerable aquatic habitat includes diving, sleeping or foraging in or thought and effort toward its evolution, context, correlates, under water, or running over the water surface. and patterns (Ydenberg and Dill 1986; Lima and Dill Accordingly, many lizards have evolved adaptations for 1990; Lima 1993; O’Steen et al. 2002; Cooper and their aquatic existence, including morphological, Frederick 2007). One way to effectively escape predators physiological, and behavioral traits (Dawson et al. 1977; is to rapidly change microhabitats (e.g., move to thicker Courtice 1985; Hsieh and Lauder 2004; Schwenk 2008; cover; Lima and Dill 1990; Sih 1997; Cooper 1998). Even but see Leal et al. 2002). more effective is to rapidly move to where a predator Adaptations in aquatic lizards are often straightforward; cannot, or will not, follow (e.g., a mouse retreating into a laterally compressed tails facilitate swimming, fringed toes small burrow when chased by a fox). Some semi-aquatic aid in running over the water surface, and diving evades species use water bodies as escape microhabitat (e.g., predators. However, it is often unclear why a particular frogs, mudskippers, snakes, crocodilians, ducks, and species is aquatic; that is, what made the species invade the beavers), and there is evidence that their would-be aquatic habitat in the first place? This question is more predators often do not follow (Carpenter 1952; Gregory difficult to answer in species that both forage underwater 1979). and use aquatic escape because these species may have Although lizard species richness is highest in arid regions invaded the aquatic environment to access aquatic food, or (Pianka and Vitt 2003), some lizard species have invaded to escape predators, or both. aquatic habitats. In a review, Bauer and Jackman (2008) A few lizard species that do not forage underwater use Copyright © 2014. Sean Doody. All Rights Reserved. 48 Herpetological Conservation and Biology water as a refuge from predators, particularly when under or in burrows or caves (Thompson 1993; Retallick and immediate threat. For example, the Australian Water Hero 1994; Anthony and Telford 1996; Turner 1999). Dragon, Physignathus lesueurii, is seldom far from water, We searched published journal articles, books, and book feeds terrestrially, and uses aquatic escape to reduce chapters, along with unpublished reports for evidence of predation risk (Barrett 1931; Anonymous 1976; Retallick predation on water dragons. These sources included and Hero 1994). However, the water dragon also sleeps dietary reviews and studies of known and suspected under water (Thompson 1993; Turner 1999; Meek and predators of Water Dragons (e.g., raptors, foxes, cats, and Avery 2008). Aquatic sleeping may also provide water rats), and anecdotal observations of predation. protection from terrestrial predators, but alternatively it Anecdotes included published notes and unpublished may confer a thermal advantage because water observations from competent herpetologists or other temperatures are generally warmer than air temperatures at biologists. We supplemented this information with four night (Turner 1999). detailed accounts of predation events, and one ‘partial Which adaptation prompted the invasion of aquatic predation’ account on Water Dragons. We then reviewed habitats in Water Dragons, the potential thermal advantage the literature on the aquatic behavior of Water Dragons, of aquatic sleeping, or predator avoidance via diving into examining the fit of these behaviors to the suite of water and/or aquatic sleeping? Although this question is predators we assembled. In this review we do not consider not directly testable, we can determine the degree to which predation of Water Dragon eggs. Water Dragons optimally exploit the aquatic medium by studying their thermoregulation (aquatic sleeping), or we RESULTS can examine the fit of their behaviors (aquatic escape and aquatic sleeping) to predation risk associated with their An exhaustive review of the literature revealed 25 known known (extant) predators. Identifying actual and potential predators of Water Dragons (Table 1). The list was predators is critical to interpreting animal behavior (Lima dominated by 11 species of birds, seven species of snakes, and Dill 1990). Such information is often difficult to and three lizard species. Observations of predation by obtain for reptiles due to the paucity of direct observations mammals (two species) and fish (two species) were rare. of predation. However, the opportunity exists to determine predominated; whereas, aquatic predators were rare (8%). the suite of predators by reviewing predation anecdotes At about 1230 on 8 January 2007, we observed a 1.0–1.2- and dietary studies of potential predators. m long Eastern Brown Snake (Pseudonaja textilis) with an In this paper we address the predation risk hypothesis by adult Water Dragon in its mouth, in the lower reaches of reviewing the known predators of water dragons through the Cotter River, Australian Capital Territory, Australia examination of published dietary records of suspected (35°20′45.94″S, 148°53′22.36″E). The snake was in the predators (e.g., raptors) and by gleaning published and stream about 1 m from the bank and clinging to a small unpublished observations of predation events. We include branch attached to the shore. The Water Dragon was up to detailed descriptions of predation events in a few cases to its shoulders in the mouth of the snake and was kicking provide insights into predator avoidance and predator and lashing its tail. After a few minutes the snake hunting tactics. We discuss how well the suite of detached from the branch and drifted downstream with the identified and suspected predators matches the antipredator dragon, seeking purchase with its tail. We are unsure if the behaviors of Water Dragons, to address why they may snake successfully swallowed the dragon. At 1025 on 24 have invaded their aquatic habitat. December 2007, we observed a 1.3-m male Eastern Brown Snake with an adult female Water Dragon in its mouth at MATERIALS AND METHODS the Australian National Botanic Gardens (ANBG), Canberra, Australian Capital Territory, Australia The Australian Water Dragon is a large, semi-aquatic (35°16′44.19″S, 149°06′37.74″E; Fig. 1). The snake held agamid lizard inhabiting riparian
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