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Komodo Dragon (Varanus komodoensis) Fact Sheet: Bibliography & Resources

Komodo Dragon (Varanus komodoensis)


Arbuckle K. 2009. Ecological function of venom in Varanus, with a compilation of dietary records from the literature. Biawak. 3(2):46–56.
Ariefiandy A, Purwandana D, Seno A, Ciofi C, Jessop TS. 2013. Can camera traps monitor Komodo dragons, a large ectothermic predator? PLoS ONE. 8(3):e58800. doi:10.1371/journal.pone.0058800.
Ariefiandy A, Purwandana D, Seno A, Chrismiawati M, Ciofi C, Jessop TS. 2014. Evaluation of three field monitoring-density estimation protocols and their relevance to Komodo dragon conservation. Biodivers Conserv. 23(10):2473–2490. doi:10.1007/s10531-014-0733-3.

Ariefiandy A, Purwandana D, Nasu SA, Surahman M, Ciofi C and Jessop T. 2015. First record of Komodo dragon nesting activity and hatchling emergence from north Flores, eastern Indonesia. Biawak. 9(1):33–35. [PDF]

Ariefiandy A, Purwandana D, Natali C, Imansyah MJ, Surahman M, Jessop TS, Ciofi C. 2015. Conservation of Komodo dragons Varanus komodoensis in the Wae Wuul nature reserve, Flores, Indonesia: a multidisciplinary approach. International Zoo Yearbook 49: 67-80.

Ariefiandy A, Purwandana D, Azmi M, Nasu SA, Mardani J, Ciofi C, Jessop TS. 2021. Human activities associated with reduced Komodo dragon habitat use and range loss on Flores. Biodivers Conserv. 30(2):461–479. doi:10.1007/s10531-020-02100-8.

Auffenberg W. 1978. Social and feeding behavior in Varanus komodoensis. In: Greenberg, N. and P. MacLean (eds.), Behavior and neurology of lizards. U.S. Department of Health, Education and Welfare, National Institute of Mental Health, pp. 301-331.

Auffenberg W.1980. The herpetofauna of Komodo, with notes on adjacent areas. Bulletin of the Florida State Museum Biological Sciences, 25(2):156.

Auffenberg W.  1981. The behavioral ecology of the Komodo monitor.  Gainesville (FL): University of Florida Press; 406 pp.

AZA. 2023. Species Survival Statistics [data table report]. Chicago (IL): Lincoln Park Zoo/AZA Population Management Center; [accessed 2023 Apr 18].

Barnard TP. 2011. Protecting the dragon: Dutch attempts at limiting access to Komodo lizards in the 1920s and 1930s. Indonesia.(92):97-123. doi:10.5728/indonesia.92.0097.

Behler JL. 1988. The Komodo monitor consortium - 1988. AAZPA, Annual Conference Proceedings,  pp. 159-161.

Birchard GF, Walsh T, Rosscoe R, and Reiben CL.1995. Oxygen uptake by Komodo dragon (Varanus komodoensis) eggs: the energetics of prolonged development in a reptile. Physiological Zoology, 68(4):622-633.

Ciofi C. 1999. The Komodo dragon. Scientific American 280(3):84-91. [Distribution map was obtained online]

Ciofi C and De Boer ME. 2004. Distribution and conservation of the Komodo monitor (Varanus komodoensis). Herpetological Journal 14(2): 99-107. 

Ciofi C. 2004. Varanus komodoensis. In: Pianka E. and D. King (eds), Varanoid lizards of the world. Bloomington (IN): Indiana University Press;. pp.197-204.

Claudino-Sales V. 2019. Komodo National Park, Indonesia. In: Coastal World Heritage Sites (Finkl CW, series ed.). Dordrecht (Netherlands): Springer Nature; p. 551-556.

D’Amore DC, Moreno K, McHenry CR, Wroe S. 2011. The effects of biting and pulling on the forces generated during feeding in the Komodo dragon (Varanus komodoensis). PLoS ONE. 6(10):e26226. doi:10.1371/journal.pone.0026226

De Lisle HF. 1996. The natural history of monitor lizards.  Malibar (FL): Kreiger Publishing Company.

Dobson JS, Zdenek CN, Hay C, Violette A, Fourmy R, Cochran C, Fry BG. 2019. Varanid lizard venoms disrupt the clotting ability of human fibrinogen through destructive cleavage. Toxins. 11(5):255. doi:10.3390/toxins11050255

Fitch AJ, Goodman AE, Donnellan SC. 2006. A molecular phylogeny of the Australian monitor lizards (Squamata: Varanidae) inferred from mitochondrial DNA sequences. Australian Journal of Zoology 54:253-269. doi:10.1071/ZO05038

Forth G. 2010. Folk knowledge and distribution of the Komodo dragon (Varanus komodoensis) on Flores Island. Journal of Ethnobiology. 30(2):289–307. doi:10.2993/0278-0771-30.2.289.

Fry BG, Vidal N, Norman IA, Vonk FI, Scheib H, Ramjan SFR, Kuruppu, Fung SK, Hedges SB, Richardson MK, Hodgson WC, Ignjatovic V, Summerhayes R, Kochva E. 2006. Early evolution of the venom system in lizards and snakes. Nature 439:584-588. doi:10.1038/nature04328.

Fry BG, Schelb H, Messenger K, Tyrell KL, Hocknull S, Wroe S, Sunagar K, Goldstein EJC, Citron DM and Jackson TNW. 2015. Poisonous snakes and Komodo dragon weaponized bacteria: which is myth and which is reality? In: Fry BG, editor. Venomous reptiles and their toxins: Evolution, Pathophysiology, and Biodiscovery. New York, NY: Oxford University Press. p. 408-413.

Gokkon B. 2019 Apr 5. Indonesia arrests 7 for allegedly selling Komodo dragons over Facebook. Mongabay.

Gray C, Marcus LC, McCarten WC, and Sappington T. 1967. Amoebiasis in the Komodo dragon, (Varanus komodoensis). International Zoo Yearbook 6:279-283.

Green B and King D. 1993. Goanna; the biology of varanid lizards, New South Wales, Australia: Wales University Press.

HAGR. 2021. Varanus komodoensis. AnAge: the animal aging and longevity database [online database]. Liverpool, UK: Human Aging and Genetic Resources [compiled by Institute of Ageing and Chronic Disease, Liverpool, UK]; [accessed 2021 May 11].

Halverson J, Spelman LH. 2002. Sex determination and its role in management. In: Komodo dragons: biology and conservation (Murphy JB, Ciofi C, de la Panouse C, Walsh T). Zoo and Aquarium Biology Conservation series.

Harlow HJ, Purwandana D, Jessop TS, Phillips JA. 2010. Size-related differences in the thermoregulatory habits of free-ranging Komodo dragons. International Journal of Zoology. 2010:1–9. doi:10.1155/2010/921371.

Hildebrandt TB, Goritz F, Pitra C, Spelman LH, Walsh T, et al. 1996.  Sonomorphological sex determination in subadult Komodo dragons. Proceedings of the American Association of Zoological Veterinarians, Puerto Vallarta, Mexico. Washington DC: Smithsonian Institution Press; p. 165-176.

Hocknull SA, Piper PJ, van den Bergh GD, Due RA, Morwood MJ, Kurniawan I. 2009. Dragon’s paradise lost: palaeobiogeography, evolution and extinction of the largest-ever terrestrial lizards (Varanidae). PLoS ONE. 4(9):e7241. doi:10.1371/journal.pone.0007241.

Holmes RB, Murray AM, Attia YS, Simons EL, Chatrath P. 2010. Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus. Palaeontology. 53(5):1099–1110. doi:10.1111/j.1475-4983.2010.00994.x.

Hudson R, Alberts A., Ellis S, Byers O (eds.). 1994. Conservation assessment and management plan for Iguanidae and Varanidae, IUCN/SSC.

Imansyah MJ, Jessop TS, Ciofi C, Akbar Z. 2008. Ontogenetic differences in the spatial ecology of immature Komodo dragons. J Zoology. 274(2):107–115. doi:10.1111/j.1469-7998.2007.00368.x.

Integrated Taxonomic Information System database (ITIS) [Internet]. [accessed 2017 Mar 21]. Varanus komodoensis.

Jessop TS, Ariefiandy A, Purwandana D, Ciofi C, Imansyah J, Benu YJ, Fordham DA, Forsyth DM, Mulder RA, Phillips BL. 2018. Exploring mechanisms and origins of reduced dispersal in island Komodo dragons. Proceedings of the Royal Society B: Biological Sciences. 285(1891):20181829. doi:10.1098/rspb.2018.1829.

Jessop T, Ariefiandy A, Azmi M, Ciofi C, Imansyah J, Purwandana D. 2021. Varanus komodoensis. The IUCN red list of threatened species [Internet]. Version 2021-2; e.T22884A123633058 [species assessed 2019 Aug 05; page accessed 2021 Sep 21]. doi:10.2305/IUCN.UK.2021-2.RLTS.T22884A123633058.en.

Jones ML. 1965. The Komodo dragon lizard, Varanus (placovaranus) komodoensis, exhibition and longevity in captivity. Unpublished manuscript. 50 pp and Appendix.

Jones AR, Jessop TS, Ariefiandy A, Brook BW, Brown SC, Ciofi C, Benu YJ, Purwandana D, Sitorus T, Wigley TML, et al. 2020. Identifying island safe havens to prevent the extinction of the World’s largest lizard from global warming. Ecol Evol. 10(19):10492–10507. doi:10.1002/ece3.6705.

Kamil PI, Susianto H, Purwandana D, Ariefiandy A. 2020. Anthropomorphic and factual approaches in Komodo dragon conservation awareness program for elementary school students: Initial study. Applied Environmental Education & Communication. 19(3):225–237. doi:10.1080/1533015X.2019.1582374.

King D, King M, Baverstock P. 1991. A new phylogeny of the Varanidae. Mertensiella 2:211-219.

Laver RJ, Purwandan D, Ariefiandy A, Imansyah J, Forsyth D, Ciofi C, Jessop TS. 2012. Life-history and spatial determinants of somatic growth dynamics in Komodo dragon populations. PLoS 7(9): e45398. doi:10.1371/journal.pone.0045398.

Lutz D, Lutz JM. 1991. Komodo: the living dragon. Salem (OR): Dimi Press.

Mendyk RW. 2015. Life expectancy and longevity of varanid lizards (Reptilia: Squamata: Varanidae) in North American zoos. Zoo Biology. 34(2):139–152. doi:10.1002/zoo.21195.

Merchant M, Henry D, Falconi R, Muscher B, Bryja J. 2013. Antibacterial activities of serum from the Komodo Dragon (Varanus komodoensis). Microbiol Res (Pavia). 4(1):4. doi:10.4081/mr.2013.e4.


Miller P, Castle K (eds.) 1995. Komodo monitor (Varanus komodoensis) population and habitat viability assessment (PHVA). Bogor, Indonesia, Briefing Book. IUCN/SSC Conservation Breeding Specialist Group: Apple Valley, MN.

Molnar R. 2004. The long and honorable history of monitors and their kin. In: Pianka E. and D. King (eds). Varanoid lizards of the world. Bloomington (IN): Indiana University Press; pp.10-67.

Murphy JB, Ciofi C, Panouse C de L, Walsh T. 2002. Komodo dragons: biology and conservation. Washington (DC): Smithsonian Institution Press.

Ouwens PA.1912. On a large Varanus species from the island of Komodo. V’s Lands Plantentuin, Plantentuin ’s Lands. 1911-1912. Bulletin du Jardin botanique de Buitenzorg. Buitenzorg [etc.].

________ Pianka E. 2004. Biogeography and phylogeny of varanoids. In: Pianka E. and D. King (eds), Varanoid Llizards of the world. Bloomington (IN): Indiana University Press. pp.68-76.

Padian K., Olsen PE. 1984. Footprints of the Komodo monitor and the trackways of fossil reptiles. Copeia 1984(3):862-871. doi:10.2307/1445147

Post TJ. 2020. Komodo Island eyes 50,000 tourists after premium membership system introduced. The Jakarta Post. [accessed 2021 Apr 13].

Post TJ., Muh. Ibnu Aqil. 2020. Komodo dragon's 'Tiananmen moment' sparks concern over prime tourism project. The Jakarta Post. [accessed 2021 Apr 6]. 

Purwandana D, Ariefiandy A, Imansyah MJ, Rudiharto H, Seno A, Ciofi C, Fordham DA, Jessop TS. 2014. Demographic status of Komodo dragons populations in Komodo National Park. Biological Conservation 171: 29-35. doi:10.1016/j.biocon.2014.01.017.

Purwandana D, Ariefiandy A, Imansyah MJ, Seno A, Ciofi C, Letnic M, Jessop TS. 2016. Ecological allometries and niche use dynamics across Komodo dragon ontogeny. Sci Nat. 103(3–4):27. doi:10.1007/s00114-016-1351-6.

Purwandana D, Imansyah MJ, Ariefiandy A, Rudiharto H, Ciofi C, Jessop TS. 2020. Insights into the nesting ecology and annual hatchling production of the Komodo dragon. Copeia. 108(4):855–862. doi:10.1643/CH-19-337.

Seal US, Manansang J, Siswomartono D, Suhartono T, Sugarjito J. (eds.).1995. Komodo monitor (Varanus komodoensis) population and habitat viability assessment workshop (PHVA). December 4-7, 1995. Taman Safari Indonesian Proceedings Cisuara, Indonesia.

Shaw C. 1963. Dragons big and bold. Oct 1963. ZOONOOZ, Vol. XXXVI (No. 10); p. 3-7.

Shine R, Somaweera R. 2019. Last lizard standing: the enigmatic persistence of the Komodo dragon. Global Ecology and Conservation. 18:e00624. doi:10.1016/j.gecco.2019.e00624.

Spelman L., Cambre RC, Walsh T,. Rosscoe R. 1996. Anesthetic techniques in Komodo dragons. Proceedings of the American Association of Zoolgical Veterinarians, Puerto Vallarta, Mexico.

Sulandari S, Zein MSA, Arida EA, Hamidy A. 2014. Molecular sex determination of captive Komodo dragons (Varanus komodoensis) at Gembira Loka Zoo, Surabaya Zoo, and Ragunan Zoo, Indonesia. HAYATI Journal of Biosciences. 21(2):65–75. doi:10.4308/hjb.21.2.65.

Sunkar A, Kusrini MD, Ramadhani FS. 2020. Role of culture in the emotional response towards komodo dragon in Komodo and Rinca Islands of Komodo National Park. Setiawan Achmadi A, Anita S, Rusdianto, Dwijayanti E, Setyo Budi A, Murniati DC, Nurinsiyah AS, Qodri A, Encilia, Elang N.H. A, editors. BIO Web Conf. 19:00021. doi:10.1051/bioconf/20201900021.

Sunter G. 2008. Management and reproduction of the Komodo dragon Varanus komodoensis Ouwens 1912 at ZSL London Zoo. International Zoo Yearbook. 42(1):172–182. doi:10.1111/j.1748-1090.2007.00029.x.

Trivedi B. (2003). Trapping Komodo dragons for conservation. National Geographic Today, January 29, 2003

UNEP. 2018. Species+: Varanus komodoensis [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2018 Nov 13].

UNESCO. 2019. Komodo National Park [Internet]. [accessed 2019 Apr 10]. Paris: United Nations Educational, Scientific and Cultural Organization.

van Hoek ML, Prickett MD, Settlage RE, Kang L, Michalak P, Vliet KA, Bishop BM. 2019. The Komodo dragon (Varanus komodoensis) genome and identification of innate immunity genes and clusters. BMC Genomics. 20(1):684. doi:10.1186/s12864-019-6029-y.

Walsh T, Rosscoe R, Birchard GF. 1993. Dragon Tales: The history, husbandry and breeding of Komodo monitors at the National Zoological Park. Vivarium, 4(6):23-26.

World Conservation Monitoring Centre. 1996. Varanus komodoensis. The IUCN red list of threatened species [Internet]. Version 2018-1; e.T22884A9396736 [species assessed 1996 Aug 01; page accessed 2018 Nov 13].

ZIMS (Zoological Information Management System). 2021. [updated 2021 Apr 19; accessed 2021 May 11].

Zug GR. 1993. Herpetology. an introductory biology of amphibians and reptiles. San Diego (CA): Academic Press Inc. [PDF]

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