Monitor lizards are large lizards in the genus Varanus. They are native to Africa, Asia, and Oceania, and one species is also found in the Americas as an invasive species. About 80 species are recognized.
Temporal range: Miocene to present
|Lace monitor (Varanus varius)|
|Combined native range of all the monitor lizards|
Monitor lizards have long necks, powerful tails and claws, and well-developed limbs. The adult length of extant species ranges from 20 cm (7.9 in) in some species, to over 3 m (10 ft) in the case of the Komodo dragon, though the extinct varanid known as megalania (Varanus priscus) may have been capable of reaching lengths more than 7 m (23 ft). Most monitor species are terrestrial, but arboreal and semiaquatic monitors are also known. While most monitor lizards are carnivorous, eating eggs, smaller reptiles, fish, birds, insects, and small mammals, some also eat fruit and vegetation, depending on where they live.
The various species cover a vast area, occurring through Africa, the Indian subcontinent, to China, the Ryukyu Islands in southern Japan, south to Southeast Asia to Thailand, Malaysia, Brunei, Indonesia, the Philippines, New Guinea, Australia, and islands of the Indian Ocean and the South China Sea. The West African Nile monitor (Varanus stellatus) is now found in South Florida and in Singapore. Monitor lizards also occurred in Europe in the Neogene, with the last known remains dating to the Middle Pleistocene.
Habits and dietEdit
Most monitor lizards are almost entirely carnivorous, consuming prey as varied as insects, crustaceans, arachnids, myriapods, mollusks, fish, amphibians, reptiles, birds, and mammals. Most species feed on invertebrates as juveniles and shift to feeding on vertebrates as adults. Deer make up about 50% of the diet of adults of the largest species, Varanus komodoensis. In contrast, three arboreal species from the Philippines, Varanus bitatawa, Varanus mabitang, and Varanus olivaceus, are primarily fruit eaters. Although normally solitary, groups as large as 25 individual monitor lizards are common in ecosystems that have limited water resources.
The genus Varanus is considered unique among animals in that its members are relatively morphologically conservative, yet show a very large size range. Finer morphological features such as the shape of the skull and limbs do vary, though, and are strongly related to the ecology of each species.
Monitor lizards maintain large territories and employ active-pursuit hunting techniques that are reminiscent of similar-sized mammals. The active nature of monitor lizards has led to numerous studies on the metabolic capacities of these lizards. The general consensus is that monitor lizards have the highest standard metabolic rates of all extant reptiles.
Monitor lizards have a high aerobic scope that is afforded, in part, by their heart anatomy. Whereas most reptiles are considered to have three-chambered hearts, the hearts of monitor lizards – as with those of boas and pythons – have a well developed ventricular septum that completely separates the pulmonary and systemic sides of the circulatory system during systole. This allows monitor lizards to create mammalian-equivalent pressure differentials between the pulmonary and systemic circuits, which in turn ensure that oxygenated blood is quickly distributed to the body without also flooding the lungs with high-pressure blood.
Anatomical and molecular studies indicate that all varanids (and possibly all lizards) are partially venomous. The venom of monitor lizards is diverse and complex, as a result of the diverse ecological niches monitor lizards occupy. Monitor lizards are oviparous, laying from seven to 37 eggs, which they often cover with soil or protect in a hollow tree stump. Some monitor lizards, including the Komodo dragon, are capable of parthenogenesis.
The family Varanidae probably originated in Asia at least 65 million years ago, although some estimates are as early as the late Mesozoic (112 million years ago). Monitor lizards probably expanded their geographic range into Africa between 49 and 33 million years ago, possibly via Iran, and to Australia and the Indonesian archipelago between 39 and 26 million years ago.
During the Late Cretaceous Period, monitor lizards or their close relatives are believed to have evolved into amphibious and then fully marine forms, the mosasaurs, some of which reached lengths of 12 m (39 ft) or more.
Snakes were believed to be more closely related to monitor lizards than any other type of extant reptile; however, snakes have been more recently proposed to be the sister group of the clade of iguanians and anguimorphs. Like snakes, monitor lizards have forked tongues, which they use to sense odors.
The genus Varanus first emerged in Laurasia. During the late Oligocene to early Miocene, the group had dispersed to Australia, New Guinea, and the Solomon Islands three separate times. By the late Miocene, the genus was also present in Africa, Arabia, Asia, and Eastern Europe.
By the Pleistocene Epoch, giant monitor lizards lived in Southeast Asia and Australasia, the best known fossil species being the megalania (Varanus priscus, a giant goanna formerly known as Megalania prisca). This species is an iconic member of the Pleistocene megafauna of Australia, thought to have survived until around 50,000 years ago.
V. doreanus species complex (V. doreanus, V. finschi, V. semotus, V. yuwonoi)
V. gouldii species complex (V. gouldii, V. rosenbergi, V. panoptes)
V. bengalensis species complex (V. bengalensis, V. nebulosus)
V. acanthurus species complex (V. acanthurus, V. baritji, V. primordius, V. storri)
V. exanthematicus species complex (V. exanthematicus, V. albigularis, V. yemenensis)
V. timorensis species complex (V. timorensis, V. auffenbergi, V. scalaris, V. similis, V. tristis)
V. niloticus species complex (V. niloticus, V. stellatus)
The tree monitors of the V. prasinus species complex (V. prasinus, V. beccarii, V. boehmei, V. bogerti, V. keithhornei, V. kordensis, V. macraei, V. reisingeri, V. telenesetes) were once in the subgenus Euprepriosaurus, but as of 2016, form their own subgenus Hapturosaurus.
V. jobiensis was once considered to be a member of the V. indicus species complex, but is now considered to represent its own species complex.
In English, they are known as "monitors" or "monitor lizards". The earlier term "monitory lizard" became rare by about 1920. The name may have been suggested by the occasional habit of varanids to stand on their two hind legs and to appear to "monitor", or perhaps from their supposed habit of "warning persons of the approach of venomous animals".
In Austronesia, where varanids are common, they are known under a large number of local names. They are usually known as biawak (Malay and Indonesian), bayawak (Filipino), binjawak or minjawak or nyambik (Javanese), or variations thereof. Other names include hokai (Solomon Islands); bwo, puo, or soa (Maluku); halo (Cebu); galuf or kaluf (Micronesia and the Caroline Islands); batua or butaan (Luzon); alu (Bali); hora or ghora (Komodo group of islands); phut (Burmese); and guibang (Manobo).
In South Asia, they are known as hangkok in Meitei, ghorpad घोरपड in Marathi, udumbu உடும்பு in Tamil and Malayalam, bilgoh in Bhojpuri, gohi (गोहि) in Maithili, in Sinhala as කබරගොයා (kabaragoya), in Telugu as udumu (ఉడుము), in Punjabi and Magahi as गोह (goh), in Assamese as gui xaap, in Odia as ଗୋଧି (godhi), and in Bengali as গোসাপ (goshaap) or গুইসাপ (guishaap), and गोह (goh) in Hindi.
Due to confusion with the large New World lizards of the family Iguanidae, the lizards became known as "goannas" in Australia. Similarly, in South African English, they are referred to as leguaans, or likkewaans, from the Dutch term for the Iguanidae, leguanen.
Some species of varanid lizards can count; studies feeding V. albigularis varying numbers of snails showed that they can distinguish numbers up to six. V. niloticus lizards have been observed to cooperate when foraging; one varanid lures the female crocodile away from her nest, while the other opens the nest to feed on the eggs. The decoy then returns to also feed on the eggs. Komodo dragons, V. komodoensis, at the National Zoo in Washington, DC, recognize their keepers and seem to have distinct personalities.
Monitor lizards have become a staple in the reptile pet trade. The most commonly kept monitors are the savannah monitor and Ackies dwarf monitor, due to their relatively small size, low cost, and relatively calm dispositions with regular handling. Among others, black-throated monitors, Timor monitors, Asian water monitors, Nile monitors, mangrove monitors, emerald tree monitors, black tree monitors, roughneck monitors, Dumeril's monitors, peach-throated monitors, crocodile monitors, and Argus monitors have been kept in captivity.
In parts of Pakistan and southern India, as well in Northeastern India particularly Assam the different parts of monitor lizards are used for a variety of medical purposes. The flesh is eaten for the relief of rheumatic pain, abdominal fat is used as a salve for skin infections, oil and fat are used to treat hemorrhoids or chronic pain, and the oil is used as an aphrodisiac lubricant (sande ka tel).
"Large-scale exploitation" of monitor lizards is undertaken for their skins, which are described as being "of considerable utility" in the leather industry. In Papua New Guinea, monitor lizard leather is used for membranes in traditional drums (called kundu), and these lizards are referred to as kundu palai or "drum lizard" in Tok Pisin, the main Papuan trade language.
The meat of monitor lizards is eaten by some tribes in India, the Philippines, Australia, South Africa and West Africa as a supplemental meat source. Both meat and eggs are also eaten in Southeast Asia such as Vietnam and Thailand as a delicacy. The meat of monitor lizards is used in Nepal for medicinal and food purpose.
According to IUCN Red List of threatened species, most of the monitor lizards species fall in the categories of least concern, but the population is decreasing globally. All but five species of monitor lizards are classified by the Convention on International Trade in Endangered Species of Wild Fauna and Flora under Appendix II, which is loosely defined as species that are not necessarily threatened with extinction but may become so unless trade in such species is subject to strict regulation to avoid use incompatible with the survival of the species in the wild. The remaining five species – V. bengalensis, V. flavescens, V. griseus, V. komodoensis, and V. nebulosus – are classified under CITES Appendix I, which outlaws international commercial trade in the species.
The yellow monitor (V. flavescens) is protected in all countries in its range except Bhutan, Nepal, India, Pakistan, and Bangladesh.
In Tamil Nadu and all other parts of South India, catching or killing of monitor lizards is banned under the Protected Species Act.
- Species marked with † are extinct
- V. bengalensis, Bengal monitor
- V. dumerilii, Dumeril's monitor, brown roughneck monitor
- V. flavescens, golden monitor, yellow monitor, short-toed monitor
- V. nebulosus, clouded monitor
- V. rudicollis, black roughneck monitor
- V. bennetti, Bennett's long-tailed monitor
- V. caerulivirens, turquoise monitor
- V. cerambonensis, Ceram monitor
- V. colei Kei Islands monitor
- V. doreanus, blue-tailed monitor
- V. douarrha, New Ireland monitor
- V. finschi, Finsch's monitor
- V. indicus, mangrove monitor
- V. jobiensis, peach-throated monitor
- V. juxtindicus, Rennell Island monitor
- V. melinus, quince monitor
- V. lirungensis, Talaud mangrove monitor
- V. obor, sago monitor
- V. rainerguentheri Rainer Günther’s monitor
- V. semotus, Mussau Island blue-tailed monitor
- V. tsukamotoi, Mariana monitor
- V. yuwonoi black-backed mangrove monitor, tricolor monitor
- V. zugorum, silver monitor, Zug's monitor
- V. beccarii, black tree monitor
- V. boehmei, golden-spotted tree monitor
- V. bogerti, Bogert's monitor
- V. keithhornei, canopy goanna, blue-nosed tree monitor, Nesbit River monitor
- V. kordensis, Biak tree monitor
- V. macraei, blue-spotted tree monitor
- V. prasinus, emerald tree monitor
- V. reisingeri  yellow tree monitor
- V. telenesetes, mysterious tree monitor, Rossell tree monitor
- V. acanthurus, spiny-tailed monitor, ridge-tailed monitor, Ackie's dwarf monitor
- V. a. acanthurus, spiny-tailed monitor
- V. a. brachyurus, common spiny-tailed monitor
- V. a. insulanicus, island spiny-tailed monitor
- V. auffenbergi, Auffenberg's monitor, peacock monitor
- V. baritji, White's dwarf monitor, black-spotted ridge-tailed monitor, lemon-throated monitor
- V. brevicauda, short-tailed monitor
- V. bushi, Pilbara stripe-tailed monitor, Bush's monitor
- V. caudolineatus, stripe-tailed monitor
- V. eremius, rusty desert monitor, pygmy desert monitor
- V. gilleni, pygmy mulga monitor
- V. glauerti, Kimberley rock monitor
- V. glebopalma, twilight monitor, black-palmed rock monitor
- V. hamersleyensis, Hamersley Range rock monitor
- V. kingorum, Kings' rock monitor
- V. mitchelli, Mitchell's water monitor
- V. pilbarensis, Pilbara rock monitor
- V. primordius northern ridge-tailed monitor
- V. scalaris, banded tree monitor
- V. semiremex rusty monitor
- V. similis, Similis monitor, spotted tree monitor
- V. sparnus, Dampier Peninsula monitor
- V. storri, Storr's monitor
- V. s. storri, eastern Storr's monitor
- V. s. ocreatus, western Storr's monitor
- V. timorensis, Timor monitor
- V. tristis
- V. t. tristis, black-headed monitor
- V. t. orientalis, freckled monitor
- V. salvadorii, crocodile monitor
- V. bitatawa, northern Sierra Madre forest monitor, butikaw, bitatawa
- V. mabitang, Panay monitor, mabitang
- V. olivaceus, Gray's monitor, butaan
- V. albigularis, rock monitor, white-throated monitor
- V. exanthematicus, savannah monitor, Bosc's monitor
- V. niloticus, Nile monitor
- V. stellatus, West African Nile monitor
- V. ornatus, ornate monitor
- V. yemenensis, Yemen monitor
- V. griseus, desert monitor
- V. g. griseus, desert monitor, grey monitor
- V. g. caspius, Caspian monitor
- V. g. koniecznyi, Indian desert monitor, Thar desert monitor
- V. nesterovi, Nesterov’s desert monitor
- V. spinulosus, spiny-necked mangrove monitor, Solomon Islands spiny monitor
- V. bangonorum, Bangon monitor
- V. cumingi, Cuming's water monitor, yellow-headed water monitor
- V. dalubhasa, Enteng’s monitor
- V. marmoratus, marbled water monitor, Philippine water monitor
- V. nuchalis large-scaled water monitor
- V. palawanensis, Palawan water monitor
- V. rasmusseni Rasmussen's water monitor
- V. salvator, Asian water monitor
- V. s. salvator, Sri Lankan water monitor
- V. s. andamanensis, Andaman water monitor
- V. s. bivittatus, two-striped water monitor, Javan water monitor
- V. s. macromaculatus, Southeast Asian water monitor
- V. s. ziegleri, Ziegler's water monitor
- V. samarensis, Samar water monitor
- V. togianus, Togian water monitor
- V. giganteus, perentie
- V. gouldii, Gould's monitor, sand monitor, sand goanna
- V. mertensi, Mertens' monitor
- V. panoptes
- V. p. panoptes, Argus monitor
- V. p. horni, Horn's monitor
- V. p. rubidus, yellow-spotted monitor
- †V. priscus, megalania
- V. rosenbergi, Rosenberg's monitor, heath monitor
- V. spenceri, Spencer's monitor
- V. varius, lace monitor
- V. komodoensis, Komodo dragon
- "Ackie Monitor Care Sheet". Reptile Range. Retrieved 8 April 2020.
- Bauer, Aaron M. (1998). Cogger, H.G.; Zweifel, R.G. (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. pp. 157–159. ISBN 0-12-178560-2.
- Ed Yong, "Florida’s Dragon Problem", The Atlantic April 20, 2016
- L, Georgalis Georgios; Andrea, Villa; Massimo, Delfino (2017-03-04). "The last European varanid: demise and extinction of monitor lizards (Squamata, Varanidae) from Europe". Journal of Vertebrate Paleontology. 37 (2): e1301946. doi:10.1080/02724634.2017.1301946. hdl:2318/1635702. ISSN 0272-4634. S2CID 90865661.
- Pianka, Eric R.; King, Dennis R; King, Ruth Allen (2004). Varanoid Lizards of the World. Bloomington, Indiana: Indiana University Press.
- Auffenberg, Walter (1981). The Behavioral Ecology of the Komodo Monitor. University Press of Florida.
- Greene, Harry W. (1986). Diet and Arboreality in the Emerald Monitor, Varanus Prasinus, with Comments on the Study of Adaptation. Chicago: Field Museum of Natural History. OCLC 14915452. OL 7155983M.
- Welton, L. J.; Siler, C. D.; Bennett, D.; Diesmos, A.; Duya, M. R.; Dugay, R.; Rico, E. L. B.; Van Weerd, M.; Brown, R. M. (2010). "A spectacular new Philippine monitor lizard reveals a hidden biogeographic boundary and a novel flagship species for conservation". Biology Letters. 6 (5): 654–658. doi:10.1098/rsbl.2010.0119. PMC 2936141. PMID 20375042.
- Struck, U; Altenbach, AV; Gaulke, M; F, Glaw (2002). "Tracing the diet of the monitor lizard Varanus mabitang by stable isotope analyses (d15N, d13C)". Naturwissenschaften. 89 (10): 470–473. doi:10.1007/s00114-002-0361-8. PMID 12384723. S2CID 12091969.
- Pianka, E.R. (1995). "Evolution of Body Size: Varanid Lizards as a Model System" (PDF). The American Naturalist. 146 (3): 398–414. doi:10.1086/285806. S2CID 85237692.
- McCurry, Matthew R.; Mahony, Michael; Clausen, Phillip D.; Quayle, Michelle R.; Walmsley, Christopher W.; Jessop, Tim S.; Wroe, Stephen; Richards, Heather; McHenry, Colin R. (2015). "The Relationship between Cranial Structure, Biomechanical Performance and Ecological Diversity in Varanoid Lizards". PLOS ONE. 10 (6): e0130625. Bibcode:2015PLoSO..1030625M. doi:10.1371/journal.pone.0130625. PMC 4479569. PMID 26106889.
- Clemente, C. J.; Thompson, G. G.; Withers, P. C. (2009). "Evolutionary relationships of sprint speed in Australian varanid lizards". Journal of Zoology. 278 (4): 270–280. doi:10.1111/j.1469-7998.2009.00559.x.
- King, D., Green, B., Knight, F. (1999). Monitors: The Biology of Varanid Lizards. Florida. Krieger Publishing Company.
- Pianka, E.R., Vitt, L.J. (2003). Lizards: Windows to the Evolution of Diversity. California. University of California Press.
- Wood SC, Johansen K, Glass ML, Maloiy GMO (1978). "Aerobic Metabolism of the Lizard Varanus exanthematicus: Effects of Activity, Temperature, and Size". Journal of Comparative Physiology B. 127 (4): 331–336. doi:10.1007/BF00738417. S2CID 19320799.
- Wang T, Altimiras J, Klein W, Axelsson M (2003). "Ventricular Haemodynamics in Python molurus: Separation of Pulmonary and Systemic Pressures". The Journal of Experimental Biology. 206 (Pt 23): 4241–5. doi:10.1242/jeb.00681. PMID 14581594.
- Fry BG, Wroe S, Teeuwisse W, van Osch JP, Moreno K, Ingle J, McHenry C, Ferrara T, Clausen P, Scheib H, Winter KL, Greisman L, Roelants K, van der Weerd L, Clemente CJ, Giannakis E, Hodgson WC, Luz S, Martelli P, Krishnasamy K, Kochva E, Kwok HF, Scanlon D, Karas J, Citron DM, ((Goldstein EJC)), Mcnaughtan JE, Norman JA (June 2009). "A central role for venom in predation by Varanus komodoensis (Komodo dragon) and the extinct giant Varanus (Megalania) priscus". PNAS. 106 (22): 8969–8974. Bibcode:2009PNAS..106.8969F. doi:10.1073/pnas.0810883106. PMC 2690028. PMID 19451641.CS1 maint: uses authors parameter (link)
- Fry BG, Vidal N, Norman JA, Vonk FJ, Scheib H, ((Ramjan SFR)), Kuruppu S, Fung K, Hedges B, Richardson MK, Hodgson WC, Ignjatovic V, Summerhays R, Kochva E (February 2006). "Early evolution of the venom system in lizards and snakes". Nature. 439 (7076): 584–588. Bibcode:2006Natur.439..584F. doi:10.1038/nature04328. PMID 16292255. S2CID 4386245.CS1 maint: uses authors parameter (link)
- Dobson, James S.; Zdenek, Christina N.; Hay, Chris; Violette, Aude; Fourmy, Rudy; Cochran, Chip; Fry, Bryan G. (May 2019). "Varanid Lizard Venoms Disrupt the Clotting Ability of Human Fibrinogen through Destructive Cleavage". Toxins. 11 (5): 255. doi:10.3390/toxins11050255. PMC 6563220. PMID 31067768.
- Smith, Kerri. "Dragon virgin births startle zoo keepers". Nature. Retrieved 2006-12-20.
- Zheng, Yuchi; Wiens, John J. (2016). "Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species". Molecular Phylogenetics and Evolution. 94 (Pt B): 537–547. doi:10.1016/j.ympev.2015.10.009. PMID 26475614.
- Schulte, JA; Melville, J; Larson, A (2003). "Molecular phylogenetic evidence for ancient divergence of lizard taxa on either side of Wallace's Line". Proceedings of the Royal Society B: Biological Sciences. 270 (1515): 597–603. doi:10.1098/rspb.2002.2272. PMC 1691285. PMID 12769459.
- Vidal, N; Marin, J; Sassi, J; Battistuzzi, FU; Donnellan, S; Fitch, AJ; Fry, BG; Vonk, FJ; Rodriguez de la Vega, RC; Couloux, A; Hedges, SB (2012). "Molecular evidence for an Asian origin of monitor lizards followed by Tertiary dispersals to Africa and Australasia". Biology Letters. 8 (5): 853–855. doi:10.1098/rsbl.2012.0460. PMC 3441001. PMID 22809723.
- "Monitor Lizards". Nature Wildlife. BBC. Archived from the original on 2013-05-02. Retrieved 2014-03-05.
- Conrad, Jack L.; Balcarcel, Ana M.; Mehling, Carl M. (2012-08-10). "Earliest Example of a Giant Monitor Lizard (Varanus, Varanidae, Squamata)". PLOS ONE. 7 (8): e41767. Bibcode:2012PLoSO...741767C. doi:10.1371/journal.pone.0041767. ISSN 1932-6203. PMC 3416840. PMID 22900001.
- Weijola, Valter; Vahtera, Varpu; Lindqvist, Christer; Kraus, Fred (2019-07-23). "A molecular phylogeny for the Pacific monitor lizards (Varanus subgenus Euprepiosaurus) reveals a recent and rapid radiation with high levels of cryptic diversity". Zoological Journal of the Linnean Society. 186 (4): 1053–1066. doi:10.1093/zoolinnean/zlz002. ISSN 0024-4082.
- Price, Gilbert J.; Louys, Julien; Cramb, Jonathan; Feng, Yue-xing; Zhao, Jian-xin; Hocknull, Scott A.; Webb, Gregory E.; Nguyen, Ai Duc; Joannes-Boyau, Renaud (2015-10-01). "Temporal overlap of humans and giant lizards (Varanidae; Squamata) in Pleistocene Australia". Quaternary Science Reviews. 125: 98–105. Bibcode:2015QSRv..125...98P. doi:10.1016/j.quascirev.2015.08.013.
- "Recommendations of the Nomenclature Committee". 1997-03-07. Archived from the original on 2008-05-18.
- Böhme, Wolfgang (2019-11-23). "The Kei Islands Monitor Lizard (Squamata: Varanidae: Varanus: Euprepiosaurus) as a Distinct Morphological, Taxonomic, and Conservation Unit". Russian Journal of Herpetology. 26 (5): 272–280. doi:10.30906/1026-2296-2019-26-5-272-280.
- Weijola, Valter (2019-03-14). "A molecular phylogeny for the Pacific monitor lizards (Varanus subgenus Euprepiosaurus) reveals a recent and rapid radiation with high levels of cryptic diversity". Zoological Journal of the Linnean Society. 186 (4): 1053–1066. doi:10.1093/zoolinnean/zlz002.
- BÖHME, WOLFGANG (2003). "Checklist of the living lizards of the world (family Varanidae)". Zool. Verh. Leiden. 341.
- Koch, Andre (2010). "Updated checklist of the living monitor lizards of the world (Squamata: Varanidae)". Bonn Zoological Bulletin. 57: 127–136.
- Koch, Andre (2010-05-06). "Unravelling The Underestimated Diversity Of Philippine Water Monitor Lizards (Squamata: Varanus Salvator Complex), With The Description Of Two New Species And A New Subspecies". Zootaxa. 2446: 1–54. doi:10.11646/zootaxa.2446.1.1.
- Mendyk, Robert (2018). "An Annotated Bibliography of Captive Reproduction in Monitor Lizards (Varanidae: Varanus). Part III. Soterosaurus". Biawak. 12.
- Brennan, Ian G.; Lemmon, Alan R.; Lemmon, Emily Moriarty; Portik, Daniel M.; Weijola, Valter; Welton, Luke; Donnellan, Stephen C.; Keogh, J. Scott (2020-02-03). "Phylogenomics of monitor lizards and the role of competition in dictating body size disparity". bioRxiv: 2020.02.02.931188. doi:10.1101/2020.02.02.931188. S2CID 211297088.
- Robert George Sprackland (1992). Giant lizards. Neptune, NJ: T.F.H. Publications. p. 61. ISBN 0-86622-634-6.
- Google ngrams
- Horatio Nelson, Matthew Henry Barker, The Life of Nelson Revised and Illustrated. With Original Anecdotes, Notes, Etc. By the Old Sailor, 1836 p. 35
- Mark K. Bayless (2004). "The local names of Pacific monitor lizards (Sauria: Varanidae) of Oceania & Indo-Malaysia, excluding Australia" (PDF). Micronesia. 37 (1): 49–55.
- Maren Gaulke (1992). "Taxonomy and biology of Philippine water monitors (Varanus salvator)". The Philippine Journal of Science. 121 (4): 345–381.
- King, Dennis & Green, Brian. 1999. Goannas: The Biology of Varanid Lizards. University of New South Wales Press. ISBN 0-86840-456-X, p. 43.
- Pianka, E.R.; King, D.R. and King, R.A. (2004). Varanoid Lizards of the World. Indiana University Press.
- Parameswaran, K (2006). "Case series of eosinophilic meningoencephalitis from South India". Annals of Indian Academy of Neurology. 9 (4): 217–222. doi:10.4103/0972-2327.29203.
- Eating Biawak. The Malaysian Life (April 2009)
- Hashmi, M. Usman Ali; Khan, M. Zaheer; Amtyaz; Huda, Nawaz Ul. "Current Status, Distribution and Threats of Varanus Species (Varanus bengalensis & Varanus griseus) in Karachi & Thatta of Sindh" (PDF). International Journal of Fauna and Biological Studies. AkiNik Publications. 1 (1): 34–38. ISSN 2347-2677. Retrieved 12 December 2013.
- "Meats We Also Eat". The Telegraph India. 2017-01-08. Retrieved 2018-08-12.
- Side, Sonny (5 June 2019). "Eating a DINOSAUR in Asia!!! RARE Mekong Delta Food you will only find here!" – via YouTube.
- Ghimire HR, Phuyal S, Shah KB (2014). "Protected species outside the protected areas: People's attitude, threats and conservation of the Yellow Monitor (Varanus flavescens) in the Far-western Lowlands of Nepal". Journal for Nature Conservation. 22 (6): 497–503. doi:10.1016/j.jnc.2014.08.003.
- "Identification Guides for Wildlife Traded in Southeast Asia". ASEAN-WEN. 2008.
- Ghimire HR, Shah KB (2014). "Status and habitat ecology of the Yellow Monitor, Varanus flavescens, in the Southeastern part of Kanchanpur District, Nepal" (PDF). Herpetological Conservation and Biology. 9 (2): 387–393.
- Ivanov, Martin; Ruta, Marcello; Klembara, Jozef; Böhme, Madelaine (2018-07-15). "A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology" (PDF). Journal of Systematic Palaeontology. 16 (9): 767–797. doi:10.1080/14772019.2017.1355338. ISSN 1477-2019. S2CID 73543240.
- Ziegler, Thomas; Schmitz, Andreas; Koch, Andre; Böhme, Wolfgang (2007). "A review of the subgenus Euprepiosaurus of Varanus (Squamata: Varanidae): morphological and molecular phylogeny, distribution and zoogeography, with an identification key for the members of the V. indicus and the V. prasimus species groups". Zootaxa 1472: 1-28.
- Weijola, Valter; Vahtera, Varpu; Koch, André; Schmitz, Andreas; Kraus, Fred (2020). "Taxonomy of Micronesian monitors (Reptilia: Squamata: Varanus): endemic status of new species argues for caution in pursuing eradication plans". Royal Society Open Science. 7 (5): 200092. Bibcode:2020RSOS....700092W. doi:10.1098/rsos.200092. PMC 7277287. PMID 32537217.
- Weijola, Valter; Sweet, Samuel (2010). "A new melanistic species of monitor lizard (Reptilia: Squamata: Varanidae) from Sanana Island, Indonesia" (PDF). Zootaxa. 2434: 17–32. doi:10.11646/zootaxa.2434.1.2.
- Weijola, Valter; Donnellan, Stephen; Lindqvist, Christer (2016). "A new blue-tailed Monitor lizard (Reptilia, Squamata, Varanus) of the Varanus indicus group from Mussau Island, Papua New Guinea". ZooKeys (568): 129–54. doi:10.3897/zookeys.568.6872. PMC 4829673. PMID 27103877.
- Varanus keithhornei, The Reptile Database
- Varanus prasinus, The Reptile Database
- Eidenmüller, Bernd; Wicker, Rudolf (2005). "Eine weitere neue Waranart aus dem Varanus prasinus-Komplex von den Insel Misol, Indonesian ". Sauria 27 (1): 3-8. (Varanus reisingeri, new species). (in German).
- Varanus baritji, The Reptile Database
- Bucklitsch, Yannick (2016-08-17). "Scale Morphology and Micro-Structure of Monitor Lizards (Squamata: Varanidae: Varanus spp.) and their Allies: Implications for Systematics, Ecology, and Conservation". Zootaxa. 4153 (1): 1–192. doi:10.11646/zootaxa.4153.1.1. PMID 27615821.
- Brendan M. Lynch (2015-01-12). "Undercover researchers expose two new species of lizard for sale on Philippine black market". KU Today. KU | The University of Kansas.
- Koch, André; Gaulke, Maren; Böhme, Wolfgang (2010). "Unravelling the underestimated diversity of Philippine water monitor Lizards (Squamata: Varanus salvator complex), with description of two new species and a new subspecies". Zootaxa 2446: 1-54. (Varanus rasmusseni, new species, p. 28).
- Varanus rasmusseni, The Reptile Database
- Merrem B (1820). Versuchs eines Systems der Amphibien: Tentamen Systematis Amphibiorum. Marburg: J.C. Krieger. xv + 191 pp. + one plate. (Varanus, new genus, p. 58). (in German and Latin).