Literatur und Schriften


Intellagama (GRAY, 1831)

Australische Wasseragame / Water Dragons

ALIBARDI, L. (1999): Formation of large micro-ornamentations in developing scales of agamine lizards. – J. Morphol., 240 (3): 251-266.

The embryonic scales of two Australian agamine lizards (Hypsilurus spinipes and Physignatus lesueuerii) derive from the undulation of the epidermis to form dome-shaped scale anlage that later become asymmetric and produce keratinized layers. Glycogen is contained in basal and suprabasal cells of the forming outer scale surface that are destined to differentiate into b-keratin cells. The outer peridermis is very flat, but the second epidermal layer, provisionally identified as an inner peridermis, is composed of large cells that accumulate vesicular bodies and a network of coarse filaments. The sequence of epidermal layers produced beneath the inner peridermis in these agamine lizards corresponds to that of previously studied lizards, but the first subperidermal layer has characteristics of both clear (keratohyalin-like granules) and oberhautchen (dark b-keratin packets) cells. This layer is here identified as an oberhautchen since it fuses with the underlying b-keratinizing cells forming large spinulae as the entire tissue becomes syncytial so that the units appear to increase in size. These spinulae very likely represent sections of honeycomb-shaped micro-ornamentations. A mesos layer appears underneath the b-layer before hatching.

BARTLETT, R.D. & P.P. BARTLETT (1997): Anoles, Basilisks, and Water Dragons. – Barron´s Educational Series, Hauppauge, New York. 96 S.

BLAKE, E. (1985): The captive breeding of Physignathus and the use of vitamin D3 with other species of lizards. – Herptile, 10 (2): 51-58.

BRAZENOV, C.W. (1932): A lizard not previously recorded from Victoria (Physignathus). – Vict. Nat., 49: 171.

CLIFFORD, H.T. & T. HAMLEY (1982): Seed dispersal by water-dragons. – Queensland Naturalist, 23 (5-6): 49.

COBORN, J. (undatiert): The Guide to Owning a Water Dragon. – t.f.h. Publications, Neptune City. 64 S.

GLAUERT, L. (1959): Herpetol. Miscel. X: Dragon Lizards. – Western Austr. Nat., 7: 10-19.

GRIESSEL, H. (1990): Wasseragamen im Terrarium. – Die Aquar. Terrar. Z., Stuttgart, 43 (9): 534-535.

LOVERIDGE, A. (1933): New agamid lizards of the genera Amphibolurus and Physignathus from Australia. – Proc. New. Engl. Zool. Blub, 13: 69-72.

MUDDE, P. (1982): Groene leguanen, basilisken en wateragamen. – Lacerta, 40 (10/11): 218-220. (02.049)

SCHRAMM, U., RUDAT, B. & W. KÜHNEL (1989): Die Nickhaut der Wasseragame. Über besondere Funktionsformen des Epithels der Plica semilunaris conjunctivae. – Verhandlungen der anatomischen Gesellschaft, 82 (1): 381-383.

STORR, G.M. (1974): Agamid lizards of the genera Caimanops, Physignathus and Diporiphora in Western Australia and Northern Territory. – Records W. Aust. Mus., 3 (2): 121-146.

VOGEL, Z. (1969): Die Mini-Saurier. Wasseragamen aus Indien und Australien. - Aquarien Magazin, Stuttgart, 3 (10): 406-408. (1098)

WERNING, H. (2002): Wasseragamen und Segelechsen. – Natur- und Tier-Verlag, Münster. 127 S.

WERNING, H. (2004): Bibliographie der Gattungen Physignathus, Lophognathus und Hydrosaurus. – Iguana-Rundschreiben, 17 (2): 18-31.



Intellagama lesueurii GRAY, 1831

Australische Wasseragame

ALDERTON, D. (1987): Spotlight. The eastern water dragon. – Aquarist and Pondkeeper, 52 (2): 11-12.

ALIBARDI, L. (2000): Epidermal structure of normal and regenerating skin of the agamine lizard Physignathus lesueurii (McCoy, 1878) wirth emphasis on the formation of the shedding layer. – Annales des Sciences naturelles Zoologie et Biologie animale, 21 (1): 27-36.

ALTMANN, H. (1969): Zur Haltung von Physignathus lesueurii. – Die Aquar. Terrar. Z., Stuttgart, 22: 94.

AMEY, A.P., COUPER, P.J. & G.M. SHEA (2012): Intellagama lesueurii (Gray, 1831): the correct combination for the Australian Eastern Water Dragon (Sauria, Agamidae). – Zootaxa, 3390: 65-67.

ANTHONY, M. & L. TELFORD (1996): Observations of the Eastern Water dragon (Physignathus lesueurii) in the northern Wet Tropics area, with a note on an unusual defensive behaviour in a juvenile. – Chondro, 4 (1): 21-22.

AUSTRALIAN HERPETOLOGICAL SOCIETY MEMBERS (1976): Observations on the eastern water dragon Physignathus lesueurii in the natural state and in captivity. – Herpetofauna, Sydney, 8 (2): 20-22.

BADIANE, A. (2017): Predation on an eastern water dragon (Intellagama lesueurii) by a heath monitor (Varanus rosenbergi) in Royal National Park, Australia. – Herpetol. Notes, 10: 339.

BAIRD, T.A., BAIRD, T.D. & R. SHINE (2012): Aggressive transition between alternative male saocial tactics in a long-lived Australian Dragon (Physignathus lesueurii) living at high density. – PloS one, 7 (8): 1-8.

Theory predicts the evolution of alternative male social tactics when intense competition coupled with the superior competitive ability of some individuals limits access to reproductive opportunities by others. How selection has shaped alternative social tactics may be especially interesting in long-lived species where size among sexually mature males varies markedly. We conducted experimental studies on long-lived eastern Australian water dragons living where competition was intense to test the hypotheses that mature males adopt alternative social tactics that are plastic, and body condition determine resource-holding potential. Approximately one-half of mature males (N = 14) defended territories using high rates of patrol and advertisement display, whereas 16 smaller mature males having lower body condition indices utilized non-territorial social tactics. Although territorial males were larger in absolute size and head dimensions, their heads were not allometrically larger. Territorial males advertised very frequently using displays involving stereotypical movements of the head and dewlap. More aggressive displays were given infrequently during baseline social conditions, but increased during periods of social instability. Female home ranges overlapped those of several territorial and non-territorial males, but females interacted more frequently with territorial males. The extreme plasticity of social tactics in this species that are dependent on body size was confirmed by two instances when relatively large non-territorial males spontaneously evicted territory owners, and by marked shifts in tactics by non-territorial males in response to temporary experimental removals of territory owners, followed (usually) by their expulsion when original owners were reinstated. The high level of social plasticity in this population where same-sex competitors are densely concentrated in preferred habitat suggests that chronic high energetic costs of defense may select for males to cycle between territorial and non-territorial social tactics depending upon their changing energetic status and their current capacity for competition with rivals.

BARRETT, C. (1931): The Gippsland Water Lizard (Physignathus lesueurii). – Vict. Nat., 47 (10): 162-165.

BAVERSTOCK, P.R. & S.C. DONNELLAN (1990): Molecular evolution in Australian dragons and skinks:progress report. – Memoirs of the Queensland Museum, 29: 323-331.

CLARK, P., JOHNSTONE, A.C.& R. ELLISON (2001): Inclusions in the erythrocytes of eastern water dragons (Physignathus lesueurii). – Aust. Vet. J., 79 (1): 61-62.

COURTICE, G.P. (1981): Respiration in the eastern water dragon, Physignathus lesueurii (Agamidae). – Comparative Biochemistry and Physiology A Comparative Physiology, 68 (3): 429-436.

COURTICE, G.P. (1981): The effect of temperature on bimodal gas exchange and the respiratory exchange ratio in the water dragon, Physignathus lesueurii. – Comparative Biochemistry and Physiology A Comparative Physiology, 68 (3): 437-441.

COURTICE, G.P. (1981): Changes in skin perfusion in response to local changes in pCO² in a diving lizard, Physignathus lesueurii. – Comparative Biochemistry and Physiology A Comparative Physiology, 69 (4): 805-807.

COURTICE, G.P. (1981): Some aspects of diving in a semi-aquatic lizard, Physignathus lesueurii (Agamidae). – Proceedings of the Australian Physiological and Pharmacological Society, 12 (2): 178 S.

COURTICE, G.P. (1985): Effect of hypoxia on cardiac vagal action in a lizard Physignathus lesueurii, and its contribution to diving bradicardia. – In: Grigg, G., Shine, R. & H. Ehmann (eds.): Biology of Australian frogs and reptiles. – Sxurrey Beatty & Sons Pty & the Royal Zoological Society of New South Wales, Chipping Norton (NSW). 373-377.

DIECKMANN, M. & H. WERNING (2014): Poster & Porträt. Intellagama lesueurii (GRAY, 1831). – Iguana, 27 (1): 17-22.

EGERT, J. (2002): Physignathus lesueurii. Care and breeding of the Eastern Water Dragon. – Reptilia (GB), Barcelona, Nr. 20: 48-56.

FISCHER, O. (2020): Wasserdrachen. – Reptilia, Münster, 25 (3): 10-17.

FRÈRE, C.H., NUGENT, D.R., LITTLEFORD-COLQUHOUN, B. & K. STRICKLAND (2015): Intellagama lesueurii Eastern water dragon: Cannibalism. - The Herpetological Bulletin 133: 38-39.

FRANCESCHINI, V. (2001): Olfactory system histochemistry in the agamid Physignathus lesueurii. – Herpetological Review, 32 (1): 7.

FRANCESCHINI, V., LAZZARI, M. & F. CIANI (2001): Lectin-binding patterns in the olfactory system of the lizard, Physignathus lesueurii. – Journal of Morphology, 247 (1): 34-38.

FRAUCA, H. (1972): Die australische Wasseragame (Physignathus lesueurii). – Die Aquar. Terrar. Z., Stuttgart, 25 (8): 280-283.

GARDINER, R.Z., DORAN, E., STRICKLAND, K., CARPENTER-BUNDHOO, L. & C. FRERE (2014): A face in the crowd: a non-invasive and cost effective photo-identification methodology to understand the fine scale movement of Eastern Water Dragons. - PLoS ONE, 9 (5): e96992.

Ectothermic vertebrates face many challenges of thermoregulation. Many species rely on behavioral thermoregulation and move within their landscape to maintain homeostasis. Understanding the fine-scale nature of this regulation through tracking techniques can provide a better understanding of the relationships between such species and their dynamic environments. The use of animal tracking and telemetry technology has allowed the extensive collection of such data which has enabled us to better understand the ways animals move within their landscape. However, such technologies do not come without certain costs: they are generally invasive, relatively expensive, can be too heavy for small sized animals and unreliable in certain habitats. This study provides a cost-effective and non-invasive method through photo-identification, to determine fine scale movements of individuals. With our methodology, we have been able to find that male eastern water dragons (Intellagama leuseurii) have home ranges one and a half times larger than those of females. Furthermore, we found intraspecific differences in the size of home ranges depending on the time of the day. Lastly, we found that location mostly influenced females’ home ranges, but not males and discuss why this may be so. Overall, we provide valuable information regarding the ecology of the eastern water dragon, but most importantly demonstrate that non-invasive photoidentification can be successfully applied to the study of reptiles.

GRIESSEL, H. (1990): Wasseragamen im Terrarium. – Die Aquar. Terrar. Z., Stuttgart, 43 (9): 534-535.

GRIGG, G.C., DRANE, D.R. & G.P. COUIRTICE (1979): Time constants of heating and cooling in the eastern water dragon, Physignathus lesueurii and some generalizations about heating and cooling in reptiles. – Journal of Thermal Biology, 4 (1): 95-103.

HARLOW, P.S. & F.M. HARLOW (1997): Captive reproduction and longevity in the eastern water dragon (Physignathus lesueurii). – Herpetofauna, Sydney, 27 (1): 154-19.

HAY, M. (1972): The breeding of Physignathus lesueurii in captivity. – Herpetofauna, Sydney, 5 (1): 2-3.

KENT, K., CRISTESCU, R.H., PIZA-ROCA, C., LITTLEFORD-COLQUHOUN, B.L., STRICKLAND, K. & C.H. FRÈRE (2019): Maternal nesting behaviour in city dragons: a species with temperature-dependent sex determination. – J. Urb. Ecol., 2019: 1-11.

Urban environments present some of the greatest challenges to species survival. This is particularly true for species that exhibit thermally sensitive traits, such as temperature-dependent sex determination (TSD). This is because urban environments not only present species with entirely novel ecosystems, but species will also experience increased temperatures. These temperature increases may result not only in offspring mortality, but also skewed population sex ratios. To persist in cities, urban dwellers with TSD will therefore need to adjust the temperature of the nesting environment, either through phenotypic plasticity or rapid evolution through natural selection. Here, we investigate the nesting ecology of a long-lived, urban dwelling reptile, the eastern water dragon (Intellagama lesueurii), to understand how a TSD species may respond to urban environments. Based on data collected from 72 nests over 2 nesting seasons, we show that city dragons not only dug significantly deeper nests than previously observed across their natural riparian habitat, but also nested in novel substrates. Furthermore, we observed a behaviour not previously described in this species, where mothers travel outside of their core home range to nest. This excursion behaviour potentially represents a greater maternal investment and is linked to the selection of specific microhabitats.

KNESE, W.C. (2018): Aus Alt mach Neu – Bau eines Großterrariums für Australische Wasseragamen. – Reptilia, Münster, 23 (1): 56-62.

LANGERWERF, B. (1998): Einfluß schwankender Temperaturen auf den Schlupf bei zwei Echsenarten. – elaphe N.F., 22-24.

LONGLEY, G. (1946-47): Notes on the hatching of the eggs of the Water Dragon (Physignathus lesueurii). – Proceedings of the Royal Zoological Society of New South Wales: 29.

MEEK, R. & R.A. AVERY (2008): Basking in the Australian water dragon Physignathus lesueurii; why do alpha males not respond to operative temperatures in the same way as adults and sub-adults? – Amphibia-Reptilia, 29: 257-262.

MEEK, R., AVERY, R. & E. WEIR (2001): Physignathus lesueurii (Australian water dragon): predation on a skink (Lampropholis delicata). – Herpetological Bulletin, 76: 31-32.

MEEK, R., WEIR, E. & G. SUTCLIFFE (2001): Nest temperatures of the Water Dragon Physignathus leseurii in southeast Australia. - Herp. Bull., (76): 26-27.

PÁČ, L. (1968): Sensory nerve endings in the joint capsules of some Lacertilia (Agama stellio, Physignathus lesueuri). – Scr. Med. (Spisy lék. Fak. Purkyn. Univ.), 41: 155-161. (in Tschechisch)

RADEK, G. (1963): De Wateragaam. – Lacerta, 21: 27-28.

RETALLICK, R.W.R. & J.-M. HERO (1994): Predation of an eastern water dragon (Physignathus lesueurii) by a common brown tree snake (Boiga irregularis). – Herpetofauna, Sydney, 24 (2): 47-48.

RINGMA, J.L. & S.W. SALISBURY (2014): Aquatic locomotor kinematics of the Eastern Water Dragon (Intellagama lesueurii). – J. Herp., 48 (2): 240-248.

Quantitative studies of the axial undulatory swimming techniques used by secondarily aquatic vertebrates have been largely restricted to crocodilians. Numerous members of the suborder Lacertilia (lizards) are also known to swim using axial undulatory techniques, but how they do so has received minimal attention from the scientific community. We investigated the morphology and undulatory locomotor kinematics adopted by the Eastern Water Dragon (Intellagama lesueurii) through observation of natural swimming and filming of animals in a flume tank with a high speed camera. We found that morphological modifications associated with improved swimming ability and correlations between wave characteristics and swimming velocity are limited to the tail. The shape of dorsal spines and the reduction in the width of transverse processes of the caudal vertebrae result in a mediolaterally compressed tail instead of the typically rounded or dorsoventrally compressed tail seen in other Australian agamids. Axial undulatory swimming in I. lesueurii was found to be conceptually similar to that of crocodilians, but the relatively long and thin terminal part of the tail produces a different shaped undulatory wave. Unlike crocodilians and fishes, I. lesueurii does not use frequency moderated velocity control. Instead, changes in velocity are solely controlled by the phase speed of the propagating wave. The combined effect of these traits is comparable efficiency and performance in the water relative to that of crocodilians and an improvement relative to terrestrial lizards.

PETERSEN, J. (1982): Physignathus lesueurii. – Nordisk Herpetologisk Forening, 25 (7): 181-182.

SLAETS, K. (2009): Physignatus lesueurii, ideaal terrariumdier en index voor de opwarming van de aarde. – Terra 24 (5): 29-33.

SLAETS, K. (2017): Intellagama lesueurii, de Australische wateragame. – Lacerta, 75 (4/5): 136-151.

SMITS, J. (2003): Bij de voorplaat: Physignathus lesueurii (GRAY, 1831). – Lacerta, 61 (4): 121-123.

THOMPSON, M.B. (1993): Estimate of the population structure of the eastern water dragon, Physignathus lesueurii (Reptilia: Agamidae), along riverside habitat. – Wildlife Research, 20 (5): 613-619.

VOGEL, Z. (0000): Die Mini-Saurier. Wasseragamen aus Indien und Australien. – Aquarien Magazin. S. 406-408.

WILSON, K.J. (1974): The relationship of oxygen supply for activity to body temperature in four species of lizards. – Copeia, 1974 (4): 920-934.

ZWINENBERG, A.J. (1977): Die australische Wasseragame (Physignathus lesueurii). – Aquaria, 24 (6): 93-97.

ZWINENBERG, A.J. (1982): Die Australische Wasseragame, Physignathus lesueurii. – Die Aquar. Terrar. Z., Stuttgart, 35 (12): 473-475.

ZWINENBERG, A.J. (1983): The Australian Physignathus lesueurii. – Nordisk Herpetologisk Forening, 26 (1): 25-30.



Intellagama lesueurii lesueurii (GRAY, 1831)

Australische Wasseragame / Eastern Water Dragon

ABRAHAM, G. (1981): Das Portrait: Physignathus l. lesueurii (GRAY 1831) – Australische Wasseragame. – Sauria, Berlin, 3 (3): 3-4.

BAXTER-GILBERT, J.H. & J.L. RILEY (2015): Natural history notes: Intellagama lesuerii lesuerii (Eastern Water Dragon). Trifid tail. – Herpetol. Rev., 46 (3): 433-434.

DALY, G. (1992): Aggressive territorial behaviour of free frange water dragons (Physignathus lesueurii lesueurii). – Herpetofauna, Sydnye, 22 (1): 37.

GIDDINGS, S. (1983): Breeding water dragons (Physignathus lesueurii lesueurii and Physignathus lesueurii howitii) in captivity. – South Australian Herpetology GroupNewsletter February, 1980: 2-3.

HARDY, C.J. & C.M. HARDY (1977): Tail regeneration and other observations in a species of agamid lizard. – Australian Zool., 19 (2): 141-148.

LANGERWERF, B. (1999): Die Australische Wasseragame Physignathus lesueurii lesueurii – ein professionell zu züchtendces Terrarientier. – Reptilia, Münster, 4 (6): 32-39.

LANGERWERF, B. & P. MANTEL (1998): Breeding of Physignathus lesueurii lesueurii in outdoor terraria. – Lacerta, 56 (3): 83-89.

SMITH, J. (1979): Notes on incubation and hatching of eggs of the eastern water dragon. – Herpetofauna, Sydney, 10 (2): 12-14.


Intellagama lesueurii howittii MCCOY, 1884

Australische Wasseragame / Gippsland Water Dragon

BAIRD, T.A., BAIRD, T.D. & R. SHINE (2013): Showing red: male coloration signals same-sex rivals in an Australian Water Dragon. – Herpetologica, 69 (4): 436-44

Sexually dimorphic coloration that plays a role in social signaling during stereotypical displays is well known in lizards. Previous studies on a large Australian agamid lizard (Eastern Water Dragon, Intellagama lesueurii) documented male-biased sex differences in ventral coloration, but males were not observed to use postures that displayed their ventral surfaces. Male resource-holding potential (RHP) was determined by body size, with the largest individuals controlling territories in preferred riparian habitat. Both large males that defended territories and those that did not had fully developed ventral coloration. Together these observations suggested that male ventral coloration does not play a role in social communication in male Water Dragons. We tested this hypothesis by recording the behavior of free-ranging males during baseline social conditions and when the level of aggression was much higher because we temporarily removed and then reinstated territory owners. Male Water Dragons performed three behavior patterns that displayed their conspicuous ventral coloration when engaged in agonistic encounters with same-sex rivals, especially when social conditions among neighboring males were unstable. Rival males responded to these displays in all instances. A large, model Water Dragon male in which we manipulated ventral color elicited lower-intensity responses from rival males when the model ventral surface was red than when it was brown. This result is also consistent with the hypothesis that red coloration plays a role in signaling and perhaps in intimidation of rivals. Water Dragon displays that revealed ventral coloration were not given when males interacted with females, suggesting red coloration does not function in advertisement to potential female mates. Our results support the hypothesis that red ventral coloration in Eastern Water Dragons functions in advertisement to rival males similar to the function of red coloration in several other vertebrates.

BRAZENOR, C.W. (1932): A lizard not previously recorded from Victoria. – Vict. Nat. (Melbourne), 49: 171.

DOODY, J.S. (2013): Natural history notes: Physignathus lesueurii (Australian Water Dragon). Mutilation by ravens. – Herpetol. Rev., 44 (4): 679-680.

DOODY, J.S., GUARINO, E., HARLOW, P.S., COREY, B. & G. MURRAY (2006): Quantifying nest site choice in reptiles using hemispherical photography and gap light analysis. – Herpetol. Rev., 37 (1): 49-52.

GIDDINGS, S. (1983): Breeding water dragons (Physignathus lesueurii lesueurii and Physignathus lesueurii howitii) in captivity. – South Australian Herpetology GroupNewsletter February, 1980: 2-3.

McCOY, F. (1884): The Gippsland water lizard. – Prod. Zool. Vict., 9: 7-10.

STRICKLAND, K., GARDINER, R., SCHULTZ, A.J. & C.H. FRERE (2014): The social life of Eastern Water Dragons: sex differences, spatial overlap and genetic relatedness. – Anim. Behav., 97: 53-61.

Understanding the ways individuals socialize with each other and how they differ temporally, spatially and phylogenetically is key to unravelling the evolutionary processes that shape social evolution. Our current knowledge of social evolution in vertebrates, however, has primarily come from bird and mammalian studies. Despite being largely understudied, reptiles remain an important piece of the puzzle in our study of social evolution; they represent a major class of vertebrates and, similar to mammals and birds, many are gregarious. Increasing our understanding of sociality in reptiles is important given that it would allow for comparisons across phylogenetically distinct vertebrate classes. In this study, we investigated the social structure of the eastern water dragon, Intellagama lesueurii, and found that males and females showed both preference and avoidance for members of either sex. Furthermore, we found sex differences in the extent of individual sociability: females generally formed stronger associations with one another than any other sex class (e.g. maleemale, maleefemale). Although association patterns correlated to some extent with home range overlap, we found no evidence of a correlation with kinship. Overall, our study presents additional evidence that sociality can evolve outside the realm of kin selection.

TURNER, G. (1999): Field observations of Gippsland Water Dragons Physignathus lesueurii howitti sleeping in water. – Herpetofauna (Austr.), 29: 49–51.

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