Habitat-linked morphological variation in the Asian water monitor lizard (<em>Varanus salvator</em>) across urban, agricultural, and forest landscapes in peninsular Malaysia

Authors

  • Ariff Azlan Alymann School of Biological Sciences, Universiti Sains Malaysia
  • Ong Song-Quan Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah
  • Muhammad Syafiq Mohd Zaludin School of Biological Sciences, Universiti Sains Malaysia
  • Mohd Uzair Rusli Sea Turtle Research Unit (SEATRU), Institute of Oceanography and Environment (INOS), Universiti Malaysia Terengganu
  • Sergio Guerrero-Sanchez Centre for Applied One Health Research and Policy Advice, Jockey Club College of Veterinary Medicine and Life Sciences, City University of Hong Kong
  • Shakinah Ravindran School of Biological Sciences, Universiti Sains Malaysia
  • Abu Hassan Ahmad School of Biological Sciences, Universiti Sains Malaysia
  • Hasber Salim School of Biological Sciences, Universiti Sains Malaysia

DOI:

https://doi.org/10.11160/bah.306

Keywords:

Asian water monitor lizard, disturbance, habitat, morphology, varanids

Abstract

The Asian water monitor lizard, Varanus salvator is a widespread carnivorous reptile in Malaysia, yet the effects of habitat disturbance on its morphology remain poorly understood. This study examined 168 individuals from urban, agricultural, and forest habitats using morphometric measurements of snout–vent length (SVL), weight, skull length, thigh width, and body condition. To assess shape-level differences independent of body size, size-corrected traits were calculated as residuals from log–log regressions against SVL. Principal Component Analysis (PCA), Linear Discriminant Analysis (LDA), and one-way MANOVA were used to evaluate morphological divergence among habitats. Results showed that morphological variation was structured primarily by habitat type rather than site-specific effects. SVL did not differ significantly across habitats, indicating similar growth potential across the disturbance gradient. However, urban lizards exhibited consistently larger relative traits, and better body condition than those from agricultural habitats, with forest individuals showing mixed patterns but closer to agricultural lizards. Lizards from agricultural habitats had the narrowest thighs and poorest condition, potentially a result of habitat specific stressors. These findings suggest that habitat type influences energy allocation to structural and performance-related traits, rather than limiting size. While V. salvator appears morphologically adaptable to anthropogenic environments, future studies incorporating sex, diet, and predator interactions are needed to clarify the drivers of trait variation and assess potential fitness consequences in disturbed landscapes.

References

Albuquerque, R.L.; Zani, P.A. & Garland, T. (2023). Lower-level predictors and behavioural correlates of maximal aerobic capacity and sprint speed among individual lizards. Journal of Experimental Biology 226: jeb244676.

Azhar, B.; Lindenmayer, D.; Wood, J.; Fischer, J.; Manning, A.; McElhinny, C. & Zakaria, M. (2012). Contribution of illegal hunting, culling of pest species, road accidents and feral dogs to biodiversity loss in established oil-palm landscapes. Wildlife Research 40: 1-9.

Beasley, J.C.; Olson, Z.H.; Selva, N. & DeVault, T.L. (2019). Ecological functions of vertebrate scavenging, In P.P. Olea, P. Mateo-Tomás & J.A. Sánchez-Zapata (eds.) Carrion Ecology and Management. Series: Wildlife Research Monographs, Vol. 2. Springer Nature, Cham, Switzerland, pp. 125-157.

Bhatt, Y. & Mori, D. (2022). Predators of the Bengal Monitor (Varanus bengalensis). Reptiles & Amphibians 29: 264-265.

Bhattacharya, S. & Koch, A. (2018). Effects of traditional beliefs leading to conservation of water monitor lizards (Varanus salvator) and threatened marshlands in West Bengal, India. Herpetological Conservation and Biology 13: 408-414.

Butler, M.A. & Losos, J.B. (2002). Multivariate sexual dimorphism, sexual selection, and adaptation in Greater Antillean Anolis lizards. Ecological Monographs 72: 541-559.

Cordier, J. M.; Aguilar, R.; Lescano, J. N.; Leynaud, G. C.; Bonino, A.; Miloch, D.; Loyola, R. & Nori, J. (2021). A global assessment of amphibian and reptile responses to land-use changes. Biological Conservation 253: 108863.

Cortés‐Parra, C.; Calderón‐Espinosa, M.L. & Jerez, A. (2022). Claws and toepads in mainland and island Anolis (Squamata: Dactyloidae): Different adaptive radiations with intersectional morpho spatial zones. Journal of Anatomy 240: 850-866.

Cota, M. (2011). Burrows with submerged and waterfilled entrances and nocturnal retirement of Varanus salvator macromaculatus in Thailand. Biawak 5: 44-47.

Cota, M.; Chan-Ard, T. & Makchai, S. (2009). Geographical distribution and regional variation of Varanus salvator macromaculatus in Thailand. Biawak 3: 134-143.

D'Amore, D.C.; Clulow, S.; Doody, J.S.; Rhind, D. & McHenry, C.R. (2018). Claw morphometrics in monitor lizards: variable substrate and habitat use correlate to shape diversity within a predator guild. Ecology and Evolution 8: 6766-6778.

Donihue, C. (2016). Microgeographic variation in locomotor traits among lizards in a human-built environment. PeerJ 4: e1776.

Elhaik, E. (2022). Principal Component Analyses (PCA)-based findings in population genetic studies are highly biased and must be reevaluated. Scientific Reports 12: 14683.

Falvey, C.H.; Aviles-Rodriguez, K.J.; Hagey, T. J. & Winchell, K.M. (2020). The finer points of urban adaptation: intraspecific variation in lizard claw morphology. Biological Journal of the Linnean Society 131: 304-318.

Fletcher, R.J.; Orrock, J.L. & Robertson, B.A. (2012). How the type of anthropogenic change alters the consequences of ecological traps. Proceedings of the Royal Society B 279: 2546–2552.

French, S.S.; Webb, A.C.; Hudson, S.B. & Virgin, E.E. (2018). Town and country reptiles: a review of reptilian responses to urbanization. Integrative and Comparative Biology 58: 948-966.

Gaulke, M. & Horn, H.G. (2004). Varanus salvator (Nominate form), In E. Pianka, D. King & R. King (eds.) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana, USA, pp. 244-257.

Gibbon, J.H.; Lim, J.; Gonzales-Abarzua, M.; Wengel, O.T.; Tsan-Yuk Lam, T. & Goossens, B. (2024). Active forest nesting site, mating and cannibalistic oophagy behaviour in the Asian water monitor Varanus salvator. The Herpetological Bulletin 168: 8-11.

Green, A.J. (2001). Mass/length residuals: measures of body condition or generators of spurious results? Ecology 82: 1473-1483.

Guerrero-Sanchez, S.; Frias, L.; Saimin, S.; Orozco-terWengel, P. & Goossens, B. (2023). The fast-food effect: costs of being a generalist in a human-dominated landscape. Conservation Physiology 11: coad055.

Guerrero-Sanchez, S.; Goossens, B.; Saimin, S. & Orozco-terWengel, P. (2021). The critical role of natural forest as refugium for generalist species in oil palm-dominated landscapes. PLoS ONE 16: e0257814.

Guerrero‐Sanchez, S.; Majewski, K.; Orozco‐terWengel, P.; Saimin, S. & Goossens, B. (2022). The effect of oil palm‐dominated landscapes on the home range and distribution of a generalist species, the Asian water monitor. Ecology and Evolution 12: e8531.

Higham, T.E.; Korchari, P.G. & McBrayer, L.D. (2011). How muscles define maximum running performance in lizards: an analysis using swing-and stance-phase muscles. Journal of Experimental Biology 214: 1685-1691.

Holzner, A.; Balasubramaniam, K.N.; Weiß, B. M.; Ruppert, N. & Widdig, A. (2021). Oil palm cultivation critically affects sociality in a threatened Malaysian primate. Scientific Reports 11: 10353.

Hutton, P. & McGraw, K.J. (2016). Urban–rural differences in eye, bill, and skull allometry in house finches (Haemorhous mexicanus). Integrative and Comparative Biology 56: 1215-1224.

IBM Corp. (2019). IBM SPSS Statistics for Windows (Version 26) [Software]. IBM Corp.

Jessop, T.S.; Smissen, P.; Scheelings, F. & Dempster, T. (2012). Demographic and phenotypic effects of human mediated trophic subsidy on a large Australian lizard (Varanus varius): meal ticket or last supper? PLoS ONE 7: e34069.

Johnson, C.A.; Sutherland, G.D.; Neave, E.; Leblond, M.; Kirby, P.; Superbie, C. & McLoughlin, P.D. (2020). Science to inform policy: linking population dynamics to habitat for a threatened species in Canada. Journal of Applied Ecology 57: 1314-1327.

Khadiejah, S.; Razak, N.; Ward-Fear, G.; Shine, R. & Natusch, D.J.D. (2019). Asian Water Monitors (Varanus salvator) remain common in Peninsular Malaysia, despite intense harvesting. Wildlife Research 46: 265-275.

Klecka, J. & Boukal, D.S. (2013). Foraging and vulnerability traits modify predator–prey body mass allometry: freshwater macroinvertebrates as a case study. Journal of Animal Ecology 82: 1031-1041.

Koch, A. & Böhme, W. (2010). Heading east: a new subspecies of Varanus salvator from Obi Island, Maluku Province, Indonesia, with a discussion about the easternmost natural occurrence of Southeast Asian water monitor lizards. Russian Journal of Herpetology 17: 299-309.

Koch, A.; Auliya, M.; Schmitz, A.; Kuch, U. & Böhme, W. (2007). Morphological studies on the systematics of South East Asian water monitors (Varanus salvator complex): nominotypic populations and taxonomic overview. Mertensiella 16: 109-180.

Kulabtong, S. & Mahaprom, R. (2014). Observation on food items of Asian water monitor, Varanus salvator (Laurenti, 1768) (Squamata Varanidae), in urban eco-system, Central Thailand. Biodiversity Journal 6: 695-698.

Luna, Á. & Rausell-Moreno, A. (2024). Unveiling the urban colonization of the Asian water monitor (Varanus salvator) across its distribution range using citizen science. PeerJ 12: e17357.

Macdonald, K.J., Doherty, T.S., Hradsky, B.A. & Driscoll, D.A. (2024). Disturbance and ecosystem management interact to shape reptile body condition. Animal Conservation 28: 303-314.

Marsh, R.L. & Bennett, A.F. (1985). Thermal dependence of isotonic contractile properties of skeletal muscle and sprint performance of the lizard Dipsosaurus dorsalis. Journal of Comparative Physiology B 155: 541-551.

Marsh, R.L. & Bennett, A.F. (1986). Thermal dependence of sprint performance of the lizard Sceloporus occidentalis. Journal of Experimental Biology 126: 79-87.

Meyers, J.J.; Nishikawa, K.C. & Herrel, A. (2018). The evolution of bite force in horned lizards: the influence of dietary specialization. Journal of Anatomy 232: 214-226.

Munstermann, M.J.; Heim, N.A.; McCauley, D.J.; Payne, J.L.; Upham, N.S.; Wang, S.C. & Knope, M.L. (2022). A global ecological signal of extinction risk in terrestrial vertebrates. Conservation Biology 36: e13852.

Næs, T. & Mevik, B.H. (2001). Understanding the collinearity problem in regression and discriminant analysis. Journal of Chemometrics 15: 413-426.

O’Donnell, K. & delBarco-Trillo, J. (2020). Changes in the home range sizes of terrestrial vertebrates in response to urban disturbance: a meta-analysis. Journal of Urban Ecology 6: juaa014.

Oro, D.; Genovart, M.; Tavecchia, G.; Fowler, M.S. & Martínez‐Abraín, A. (2013). Ecological and evolutionary implications of food subsidies from humans. Ecology Letters 16: 1501-1514.

Puan, C.L.; Goldizen, A.W.; Zakaria, M.; Hafidzi, M.N. & Baxter, G.S. (2011). Relationships among rat numbers, abundance of oil palm fruit and damage levels to fruit in an oil palm plantation. Integrative Zoology 6: 130-139.

Putman, B.J. & Tippie, Z.A. (2020). Big city living: a global meta-analysis reveals positive impact of urbanization on body size in lizards. Frontiers in Ecology and Evolution 8: 580745.

Quah, E.; Lwin, K.; Cota, M.; Grismer, L.; Neang, T.; Wogan, G.; McGuire, J.; Wang, L.; Rao, D.-Q.; Auliya, M. & Koch, A. (2021). Varanus salvator, In The IUCN Red List of Threatened Species 2021. International Union for Conservation of Nature, Gland, Switzerland.

Randow, H. (1932). Fauna und flora von Dehiwala auf Ceylon. Wochenschrift für Aquarien-und Terrarienkunde 29: 471-473.

Ravindran, S.; Noor, H.M. & Salim, H. (2022). Anticoagulant rodenticide use in oil palm plantations in Southeast Asia and hazard assessment to non-target animals. Ecotoxicology 31: 976-997.

Rusli, M.U.; Chen, G.N.; Booth, D.T. & Lei, J. (2020). Diet preference and activity of Asian water monitor at Chagar Hutang turtle sanctuary. Journal of Sustainability Science and Management 15: 61-67.

Šálek, M.; Drahníková, L. & Tkadlec, E. (2015). Changes in home range sizes and population densities of carnivore species along the natural to urban habitat gradient. Mammal Review 45: 1-14.

Sebastián-González, E.; Barbosa, J.M.; Pérez-García, J.M.; Morales-Reyes, Z.; Botella, F.; Olea, P.P.; Mateo-Tomás, P.; Moleón, M.; Hiraldo, F.; Arrondo, E.; Donázar, J.A.; Cortés-Avizanda, A.; Selva, N.; Lambertucci, S.A.; Bhattacharjee, A.; Anadón, J.D.; Abernethy, E.; Rhodes, O.E.Jr.; Turner, K., ... & Sánchez-Zapata, J.A. (2019). Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale. Global Change Biology 25: 3005-3017.

Shine, R.; Ambariyanto; Harlow, P.S. & Mumpuni (1998). Ecological traits of commercially harvested water monitors, Varanus salvator, in northern Sumatra. Wildlife Research 25: 437-447.

Shine, R.; Harlow, P.S. & Keogh, J.S. (1996). Commercial harvesting of giant lizards: the biology of Water Monitors Varanus salvator in southern Sumatra. Biological Conservation 77: 125-134.

Shochat, E.; Warren, P.S.; Faeth, S.H.; McIntyre, N.E. & Hope, D. (2006). From patterns to emerging processes in mechanistic urban ecology. Trends in Ecology & Evolution 21: 186-191.

Sion, G.; Watson, M.J. & Bouskila, A. (2021). Measuring body condition of lizards: a comparison between non-invasive dual-energy X-ray absorptiometry, chemical fat extraction and calculated indices. Frontiers in Zoology 18: 1-9.

Smith, J.G.; Brook, B.W.; Griffiths, A.D. & Thompson, G.G. (2007). Can morphometrics predict sex in Varanids? Journal of Herpetology 41: 133-140.

Stanner, M. (2020). A case of arboreality in an adult water monitor (Varanus salvator macromaculatus). Biawak 14: 56-57.

Taverne, M.; King-Gillies, N.; Krajnović, M.; Lisičić, D.; Mira, Ó.; Petricioli, D.; Sabolić, I.; Štambuk, A.; Tadić, Z.; Vigliotti, C.; Wehrle, B. & Herrel, A. (2020). Proximate and ultimate drivers of variation in bite force in the insular lizards Podarcis melisellensis and Podarcis sicula. Biological Journal of the Linnean Society 131: 88-108.

Tee, S.L.; Samantha, L.D.; Kamarudin, N.; Akbar, Z.; Lechner, A.M.; Ashton‐Butt, A. & Azhar, B. (2018). Urban forest fragmentation impoverishes native mammalian biodiversity in the tropics. Ecology and Evolution 8: 12506-12521.

Twining, J.P.; Bernard, H. & Ewers, R.M. (2017). Increasing land-use intensity reverses the relative occupancy of two quadrupedal scavengers. PloS ONE 12: 1-13.

Urošević, A.; Ljubisavljević, K. & Ivanović, A. (2014). Variation in skull size and shape of the Common wall lizard (Podarcis muralis): allometric and non-allometric shape changes. Contributions to Zoology 83: 67-77.

Uyeda, L. (2009). Garbage appeal: relative abundance of Water Monitor Lizards (Varanus salvator) correlates with presence of human food leftovers on Tinjil Island, Indonesia. Biawak 3: 9-17.

Uyeda, L.; Iskandar, E.; Purbatrapsila, A.; Pamungkas, J.; Wirsing, A. & Kyes, R. (2014). Water Monitor Lizard (Varanus salvator) satay: A treatment for skin ailments in Muarabinuangeun and Cisiih, Indonesia. Biawak 8: 35–38.

Uyeda, L.T.; Iskandar, E.; Kyes, R.C. & Wirsing, A.J. (2015). Encounter rates, agonistic interactions, and social hierarchy among garbage-feeding water monitor lizards (Varanus salvator bivittatus) on Tinjil Island, Indonesia. Herpetological Conservation and Biology 10: 753-764.

Verwaijen, D.; Van Damme, R. & Herrel, A. (2002). Relationships between head size, bite force, prey handling efficiency and diet in two sympatric lacertid lizards. Functional Ecology 16: 842-850.

Wongtienchai, P.; Lapbenjakul, S.; Jangtarwan, K.; Areesirisuk, P.; Mahaprom, R.; Subpayakom, N.; Singchat, W.; Sillapaprayoon, S.; Muangmai, N.; Songchan, R.; Baicharoen, S.; Duengkae, P.; Peyachoknagul, S. & Srikulnath, K. (2021). Genetic management of a Water Monitor Lizard (Varanus salvator macromaculatus) population at Bang Kachao Peninsula as a consequence of urbanization with Varanus Farm Kamphaeng Saen as the first captive research establishment. Journal of Zoological Systematics and Evolutionary Research 59: 484-497.

Xanthopoulos, P.; Pardalos, P.M. & Trafalis, T.B. (2013). Linear discriminant analysis. Robust data mining, 27-33.

Young, B.A.; Dumais, J.; John, N.; Lyons, B.; Macduff, A.; Most, M.; Reiser, N.A. & Reiser, P.J. (2016). Functional segregation within the muscles of aquatic propulsion in the Asiatic water monitor (Varanus salvator). Frontiers in Physiology 7: 380.

Zemp, D.C.; Ehbrecht, M.; Seidel, D.; Ammer, C.; Craven, D.; Erkelenz, J.; Irawan, B.; Sundawati, L.; Hölscher, D. & Kreft, H. (2019). Mixed-species tree plantings enhance structural complexity in oil palm plantations. Agriculture, Ecosystems and Environment 283: 106564.

Published

2025-12-31

Issue

Section

Research Papers