Komodo Research Articles

Invasive toxic prey may imperil the survival of an iconic giant lizard, the Komodo dragon.

Although invasive species constitute a major threat to global biodiversity, the introduction of toxic prey into naïve predator faunas may become particularly destructive. An example of such an introduction was the deliberate release of the highly toxic cane toad (Bufo marinus) to the toad-free Australian continent in 1935. Naïve large Australian varanid lizards have recently been shown to suffer a massive increase in mortality (> 95%) when attempting to feed on this toxic amphibian. The high susceptibility of Australian varanids to toad toxin is caused by minor mutations in the sodiumpotassium- ATPase enzyme. In the present study we show that Komodo dragons (Varanus komodoensis) have similar mutations within this enzyme as observed in Australian varanids demonstrating that dragons are extremely susceptible to toad toxin. During the last decade the black-spined toad (Bufo melanostictus) has been able to invade areas close to the five toad-free islands constituting the habitat of Komodo dragons. An invasion of highly toxic black-spined toads into dragon habitats may therefore cause similar dramatic increase in dragon mortality as recorded in Australian varanids imperiling the long-term survival of this giant and iconic lizard.

How to train your Komodo dragon

What do you do when a giant carnivorous lizard starts heading straight for you? Try giving it a good scratch behind the ears

Comparative insights using the molecular homology model of BDNF (Brain derived neurotrophic factor) of Varanus komodoensis and the known NGF (Nerve growth factor) structure of Naja atra.

BDNF (Brain derived neurotrophic factor) is a secretion protein and a member of the neurotrophin family of growth factors. Structural and functional characterization of BDNF Varanus komodoensis is of interest while its structure remains unknown. Thus, a homology molecular model of BDNF was constructed for gleaning possible structural insights. The model was compared with the structure of the homologous NGF (Nerve growth factor, another member of neuro-trophin family) from Naja atra. Comparative structural analysis of the models showed structural similarities with their predicted cavities for the interpretation of potential functional analogy.

The origins and persistence of Homo floresiensis on Flores: biogeographical and ecological perspectives

The finding of archaeological evidence predating 1 Ma and a small hominin species (Homo floresiensis) on Flores, Indonesia, has stimulated much research on its origins and ancestry. Here we take a different approach and examine two key questions – 1) how did the ancestors of H. floresiensis reach Flores and 2) what are the possibilities for estimating the likelihood of hominin persistence for over 1 million years on a small island? With regard to the first question, on the basis of the biogeography we conclude that the mammalian, avian, and reptilian fauna on Flores arrived from a number of sources including Java, Sulawesi and Sahul. Many of the terrestrial taxa were able to float or swim (e.g. stegodons, giant tortoises and the Komodo dragon), while the rodents and hominins probably accidentally rafted from Sulawesi, following the prevailing currents. The precise route by which hominins arrived on Flores cannot at present be determined, although a route from South Asia through Indochina, Sulawesi and hence Flores is tentatively supported on the basis of zoogeography. With regards to the second question, we find the archaeological record equivocal. A basic energetics model shows that a greater number of small-bodied hominins could persist on Flores than larger-bodied hominins (whether H. floresiensis is a dwarfed species or a descendent of an early small-bodied ancestor is immaterial here), which may in part explain their apparent long-term success. Yet the frequent tsunamis and volcanic eruptions in the region would certainly have affected all the taxa on the island, and at least one turnover event is recorded, when Stegodon sondaari became extinct. The question of the likelihood of persistence may be unanswerable until we know much more about the biology of H. floresiensis.

Antibacterial activities of serum from the Komodo Dragon (Varanus komodoensis)

Komodo dragons (Varanus komodoensis) are able to feed on large prey items by injecting a dose of toxic bacteria with their bite that, over time, kills the prey by systemic infection. Dragons also suffer bites from other members of their own species during territorial disputes and feeding frenzies. However, they do not suffer the same fate as their prey, suggesting that they have developed a strong immunity to bacterial infections. This study was undertaken to determine the antibacterial activities of serum from the Komodo dragon. Bacterial cultures were treated with different volumes serum from Varanus komodoensis and the growth was monitored by optical density at 430 nm. In addition, the serum was treated with protease, chelators of divalent metal ions, or with mild heat to determine the mechanism of antibacterial activities. Treatment of bacterial cultures with serum from Komodo dragons (Varanus komodoensis) resulted in a volume-dependent decrease in bacterial growth. Cultures of Escherichia coli, Staphylococcus aureus, and Klebsiella oxytoca exhibited moderate-strong growth inhibition by V. komodoensis serum, while cultures of Streptococcus epidermitis, Salmonella typhimurium, Providencia stuartii, and Shigella flexneri were nearly completely obliterated for 24 h by only 10% (v/v) serum. The antibacterial activity of V. komodensis serum occurred very rapidly, as 18% of E. coli growth was inhibited by a five min exposure to serum. Furthermore, 10- and 20-min incubations of E. coli with serum from V. komodoensis resulted in 43 and 68% inhibition of bacterial growth, respectively. The bactericidal capacity of the serum against E. coli was 2,075,000 bacteria/μL serum, and was inhibited by mild heat treatment, pronase, EDTA, and phosphate, indicating that the anti-bacterial action is most probably due to the presence of a potent serum complement protein system.

Water in the Park by Emily Jenkins, and: Boris Gets a Lizard by Andrew Joyner, and: Boris on the Move by Andrew Joyner (review)

Reviewed by: Water in the Park by Emily Jenkins, and: Boris Gets a Lizard by Andrew Joyner, and: Boris on the Move by Andrew Joyner Jeannette Hulick Joyner, Andrew . Boris Gets a Lizard; written and illus. by Andrew Joyner. Branches/Scholastic, 2013. [80p]. Trade ed. ISBN 978-0-545-48446-6 $15.99 Paper ed. ISBN 978-0-545-48447-3 $4.99 E-book ed. ISBN 978-0-545-48783-2 $4.99 Reviewed from galleys R Gr. 1-2. Boris on the Move; written and illus. by Andrew Joyner. Branches/Scholastic, 2013. [80p]. Trade ed. ISBN 978-0-545-48442-8 $15.99 Paper ed. ISBN 978-0-545-48443-5 $4.99 E-book ed. ISBN 978-0-545-48782-5 $4.99 Reviewed from galleys Ad Gr. 1-2. In Boris on the Move, young warthog Boris dreams of adventure, and thus is thrilled when his family uproots their previously stationary RV-like bus from boring Hogg Bay and head to parts unknown. He's sadly disappointed, however, when the bus only goes as far as Greater Hogg Bay Conservation Park and a grumpy Boris has to partake in a family hike. While he doesn't encounter the lion he'd feared, he does meet a nice stray cat, who is quickly adopted and dubbed "Lion." In Boris Gets a Lizard, Boris invites the local zoo to send their Komodo dragon on vacation to Boris' home and excitedly tells all his friends about his impending visitor. When it turns out the Komodo dragon doesn't make home visits, he quickly substitutes a skink for his classmates' viewing, claiming it's a baby Komodo dragon, but it escapes, sending everyone into a panic. The plot in on the Move is lackluster and rambling, but Boris is a likable guy, and the development of Gets a Lizard is much stronger and more satisfying. While the page count is longer than that of many easy readers, the numerous full-color illustrations fill much of that space; the brief text will be decodable to many primary-graders despite some challenging vocabulary. Joyner's illustrations are crisply and tidily composed, and the gray shaggy-haired but anthropomorphized warthogs have a springy energy. Boris' desires for adventure and for exotic pets will resonate with most kids; readers who want a "thicker" book but who aren't yet quite up to the challenge of a lengthier chapter book will find these Australian imports fill the bill nicely. [End Page 514] Copyright © 2013 The Board of Trustees of the University of Illinois

ANAEROBIC AND AEROBIC BACTERIOLOGY OF THE SALIVA AND GINGIVA FROM 16 CAPTIVE KOMODO DRAGONS (VARANUS KOMODOENSIS): NEW IMPLICATIONS FOR THE “BACTERIA AS VENOM” MODEL

It has been speculated that the oral flora of the Komodo dragon (Varanus komodoensis) exerts a lethal effect on its prey; yet, scant information about their specific oral flora bacteriology, especially anaerobes, exists. Consequently, the aerobic and anaerobic oral bacteriology of 16 captive Komodo dragons (10 adults and six neonates), aged 2-17 yr for adults and 7-10 days for neonates, from three U.S. zoos were studied. Saliva and gingival samples were collected by zoo personnel, inoculated into anaerobic transport media, and delivered by courier to a reference laboratory. Samples were cultured for aerobes and anaerobes. Strains were identified by standard methods and 16S rRNA gene sequencing when required. The oral flora consisted of 39 aerobic and 21 anaerobic species, with some variation by zoo. Adult dragons grew 128 isolates, including 37 aerobic gram-negative rods (one to eight per specimen), especially Enterobacteriaceae; 50 aerobic gram-positive bacteria (two to nine per specimen), especially Staphylococcus sciuri and Enterococcusfaecalis, present in eight of 10 and nine of 10 dragons, respectively; and 41 anaerobes (one to six per specimen), especially clostridia. All hatchlings grew aerobes but none grew anaerobes. No virulent species were isolated. As with other carnivores, captive Komodo oral flora is simply reflective of the gut and skin flora of their recent meals and environment and is unlikely to cause rapid fatal infection.

Monitoring the ungulate prey of the Komodo dragonVaranus komodoensis: distance sampling or faecal counts?

Monitoring the abundances of prey is important for informing the management of threatened and endangered predators. We evaluated the usefulness of faecal counts and distance sampling for monitoring the abundances of rusa deerRusa timorensis, feral pigSus scrofaand water buffaloBubalus bubalis, the three key prey of the Komodo dragonVaranus komodoensis, at 11 sites on five islands in and around Komodo National Park, eastern Indonesia. We used species‐specific global detection functions and cluster sizes (i.e. multiple covariates distance sampling) to estimate densities of rusa deer and feral pig, but there were too few observations to estimate densities of water buffalo. Rusa deer densities varied from from 2.5 to 165.5 deer/km2with coefficients of variation (CVs) of 15‐105%. Feral pig densities varied from 0.0 to 25.2 pigs/km2with CVs of 25‐106%. There was a positive relationship between estimated faecal densities and estimated population densities for both rusa deer and feral pig: the form of the relationship was non‐linear for rusa deer, but there was similar support for linear and non‐linear relationships for feral pig. We found that faecal counts were more useful when ungulate densities were too low to estimate densities with distance sampling. Faecal count methods were also easier for field staff to conduct than distance sampling. Because spatial and temporal variation in ungulate density is likely to influence the population dynamics of the Komodo dragon, we recommend that annual monitoring of ungulates in and around Komodo National Park be undertaken using distance sampling and faecal counts. The relationships reported here will also be useful for managers establishing monitoring programmes for feral pig, rusa deer and water buffalo elsewhere in their native and exotic ranges.

Can Camera Traps Monitor Komodo Dragons a Large Ectothermic Predator?

Camera trapping has greatly enhanced population monitoring of often cryptic and low abundance apex carnivores. Effectiveness of passive infrared camera trapping, and ultimately population monitoring, relies on temperature mediated differences between the animal and its ambient environment to ensure good camera detection. In ectothermic predators such as large varanid lizards, this criterion is presumed less certain. Here we evaluated the effectiveness of camera trapping to potentially monitor the population status of the Komodo dragon (Varanus komodoensis), an apex predator, using site occupancy approaches. We compared site-specific estimates of site occupancy and detection derived using camera traps and cage traps at 181 trapping locations established across six sites on four islands within Komodo National Park, Eastern Indonesia. Detection and site occupancy at each site were estimated using eight competing models that considered site-specific variation in occupancy (ψ)and varied detection probabilities (p) according to detection method, site and survey number using a single season site occupancy modelling approach. The most parsimonious model [ψ (site), p (site*survey); ω = 0.74] suggested that site occupancy estimates differed among sites. Detection probability varied as an interaction between site and survey number. Our results indicate that overall camera traps produced similar estimates of detection and site occupancy to cage traps, irrespective of being paired, or unpaired, with cage traps. Whilst one site showed some evidence detection was affected by trapping method detection was too low to produce an accurate occupancy estimate. Overall, as camera trapping is logistically more feasible it may provide, with further validation, an alternative method for evaluating long-term site occupancy patterns in Komodo dragons, and potentially other large reptiles, aiding conservation of this species.

Life-History and Spatial Determinants of Somatic Growth Dynamics in Komodo Dragon Populations

Somatic growth patterns represent a major component of organismal fitness and may vary among sexes and populations due to genetic and environmental processes leading to profound differences in life-history and demography. This study considered the ontogenic, sex-specific and spatial dynamics of somatic growth patterns in ten populations of the world’s largest lizard the Komodo dragon (Varanus komodoensis). The growth of 400 individual Komodo dragons was measured in a capture-mark-recapture study at ten sites on four islands in eastern Indonesia, from 2002 to 2010. Generalized Additive Mixed Models (GAMMs) and information-theoretic methods were used to examine how growth rates varied with size, age and sex, and across and within islands in relation to site-specific prey availability, lizard population density and inbreeding coefficients. Growth trajectories differed significantly with size and between sexes, indicating different energy allocation tactics and overall costs associated with reproduction. This leads to disparities in maximum body sizes and longevity. Spatial variation in growth was strongly supported by a curvilinear density-dependent growth model with highest growth rates occurring at intermediate population densities. Sex-specific trade-offs in growth underpin key differences in Komodo dragon life-history including evidence for high costs of reproduction in females. Further, inverse density-dependent growth may have profound effects on individual and population level processes that influence the demography of this species.

A novel varanic acid epimer – (24R,25S)-3α,7α,12α,24-tetrahydroxy-5β-cholestan-27-oic acid – is a major biliary bile acid in two varanid lizards and the Gila monster

A key intermediate in the biosynthetic pathway by which C24 bile acids are formed from cholesterol has long been considered to be varanic acid, (24ξ,25ξ)-3α,7α,12α-24-tetrahydroxy-5β-cholestan-27-oic acid. The (24R,25R)-epimer of this tetrahydroxy bile acid, in the form of its taurine N-acyl amidate, was thought to be the major biliary bile acid in lizards of the family Varanidae. We report here that a major biliary bile acid of three lizard species – the Komodo dragon (Varanus komodoensis), Gray’s monitor (Varanus olivaceus), and the Gila monster (Heloderma suspectum) – is a novel epimer of varanic acid. The epimer was shown to be (24R,25S)-3α,7α,12α,24-tetrahydroxy-5β-cholestan-27-oic acid (present in bile as its taurine conjugate). The structure was established by mass spectroscopy and by 1H and 13C nuclear magnetic spectroscopy, as well as by synthesis of the compound.

Fundamental situations in teaching biology: The case of parthenogenesis

This theoretical paper considers the notion of fundamental situation in the sense of Brousseau’s theory of didactical situations. It introduces some precise elements of this theory in which a teacher provides an environment for student work that aims to enable students, through constructive inquiry, to acquire well defined pieces of scientific knowledge. Situations become fundamental if they not only allow, but force students to construct the target knowledge. A classical example from mathematics is presented, where the target knowledge is a theorem of plane geometry presented as a puzzle. Then a new fundamental situation in biology is described for parthenogenetic reproduction, which has recently turned out to occur in Komodo dragons. An explicit demand to generate and test hypotheses that could explain the given example of dragon reproduction, using authentic DNA data, is given to students. The paper concludes with an analysis of the extent to which this fundamental situation in biology is authentic to the theory of didactical situations.

The Role of New Media in Facilitating Entrepreneurship in Tourism - Case Study: Promoting Komodo Island in East Nusa Tenggara

Tourism is an important of the Indonesia economy as well as a significant soure of the country’s foreign exchange revenues. According to the Center of Data and Information- Ministry of Culture and Tourism, the growth of foreign visitor arrivals to Indonesia has increased rapidly by 9.61 percent since 2010 to present.
 One of the most potencial tourism destinations is Komodo Island located in East Nusa Tenggara. With the island’s unique qualities, which include of habitat of the Komodo dragons and beautiful and exotic marine life, it is likely to be one of the promising tourism destinations in Indonesia and in the world. In 1986, the island has been declarated as a World Heritage site by UNESCO.
 The Ministry of Culture and Tourism countinously promotes many of the country’s natural potential in tourism through various media: printed media, television and especially new media. However, there challenges for Indonesian tourism industry in facilitating entrepreneurship skills among the local people in East Nusa Tenggara. According to the Central Boreau of Statistics (2011), East Nusa Tenggara is considered as one of the poorest provinces in Indonesia where the economy is lower than the average, with a high inflation of 15% , and unemployment of 30%. This research is needed to explorefurther the phenomenon behind the above facts, aiming at ex-amining the role of new media in facilitating entrepreneurship in the tourism industry in Komodo Island. The result of this study are expected to provide insights that can help local tourism in East Nusa Tenggara.
 Keywords: Tourism, Enterpreneurship, New Media

Infection in a female Komodo dragon (Varanus komodoensis) caused by Mycobacterium intracellulare: a case report

In early 2002, a bean-like whitish nodule 2 × 1 × 1 cm in size was diagnosed on the tongue of a female Komodo dragon (Varanus komodoensis) kept in a zoological garden in the Czech Republic. The nodule was removed at surgery and histopathological examination revealed a specific granulomatous inflammation. The granuloma contained a necrotic mass surrounded by a variable layer of palisading epithelioid macrophages and multinucleated giant cells with a variable admixture of lymphocytes and plasma cells. Signs of mineralisation within the granuloma were not observed. Using Ziehl-Neelsen (ZN) staining the presence of acid-fast rods (AFR) was determined and the presence of mycobacteria was confirmed by PCR. Four years later, this female died after multiple injuries caused by a male during mating. Necropsy and histopathological examinations revealed granulomatous pneumonia and myocarditis with ZN-positive AFR within granulomas. Mycobacteria were cultured from 13 of 19 tissue samples: Mycobacterium intracellulare was confirmed by sequencing of isolates from multiple affected organs including the respiratory tract, tail muscle, inguinal lymph nodes and blood; Mycobacterium sp. were also isolated from the tongue. In addition, mycobacteria were detected in 15 (46.9%) of 32 environmental samples examined in both years. M. intracellulare was detected in water sediment from the female’s terrarium in 2002, and in faeces and peat from the terrarium and in water sediment from the male terrarium in 2006. Except for M. intracellulare (n = 4), M. smegmatis (n = 1), M. a. hominissuis (n = 3), M. fortuitum (n = 2), M. interjectum (n = 1), M. peregrinum/alvei/septicum (n = 1) and Mycobacterium sp. (n = 2) were also isolated from different environmental samples.    

The structural and functional diversification of the Toxicofera reptile venom system

The evolutionary origin and diversification of the reptilian venom system is described. The resolution of higher-order molecular phylogenetics has clearly established that a venom system is ancestral to snakes. The diversification of the venom system within lizards is discussed, as is the role of venom delivery in the behavioural ecology of these taxa (particularly Varanus komodoensis). The more extensive diversification of the venom system in snakes is summarised, including its loss in some clades. Finally, we discuss the contentious issue of a definition for “venom”, supporting an evolutionary definition that recognises the homology of both the venom delivery systems and the toxins themselves.

Characterization of serum complement activity in serum of the Komodo dragon (<i>Varanus komodoensis</i>)

Incubation of different volumes of serum from the Komodo dragon (Varanus komodoensis) with sheep red blood cells (SRBCs) resulted in volume-dependent hemolysis, as measured spectrophotometrically at 540 nm. The hemolysis occurred rapidly, with almost 90% of the hemolytic activity occurring within 20 min of incubation. A thermal profile showed that Komodo dragon serum exhibited low activity from 5- 20℃, but exerted maximum activity at 35℃, which was substantially reduced at 40℃. The maximum activity was observed near optimal temperatures to which Komodo dragons thermoregulate. Mild heat treatment of Komodo dragon serum (56℃, 30 min) depleted the ability to hemolyze SRBCs. In addition, preincubation of Komodo dragon serum with only 5 mM EDTA or phosphate, both chelators of divalent metal ions, reduced the hemolytic activity sharply. These results indicate that the hemolytic activity was due to the presence of a potent serum complement system. Incubation of Komodo dragon serum with 5 mM EDTA and 15 mM Ca2+ or Mg2+, but not Ba2+, Zn2+, or Fe2+, completely restored activity. These results indicate that Komodo dragon serum complement activity requires the presence of Mg2+ or Ca2+. This is the first assessment of innate immune activity of a Varanid.

Using striated tooth marks on bone to predict body size in theropod dinosaurs: a model based on feeding observations of Varanus komodoensis, the Komodo monitor

Mesozoic tooth marks on bone surfaces directly link consumers to fossil assemblage formation. Striated tooth marks are believed to form by theropod denticle contact, and attempts have been made to identify theropod consumers by comparing these striations with denticle widths of contemporaneous taxa. The purpose of this study is to test whether ziphodont theropod consumer characteristics can be accurately identified from striated tooth marks on fossil surfaces. We had three major objectives (1) to experimentally produce striated tooth marks and explain how they form; (2) to determine whether body size characteristics are reflected in denticle widths; and (3) to determine whether denticle characters are accurately transcribed onto bone surfaces in the form of striated tooth marks. We conducted controlled feeding trials with the dental analogue Varanus komodoensis (the Komodo monitor). Goat (Capra hircus) carcasses were introduced to captive, isolated individuals. Striated tooth marks were then identified, and striation width, number, and degree of convergence were recorded for each. Denticle widths and tooth/body size characters were taken from photographs and published accounts of both theropod and V. komodoensis skeletal material, and regressions were compared among and between the two groups. Striated marks tend to be regularly striated with a variable degree of branching, and may co-occur with scores. Striation morphology directly reflects contact between the mesial carina and bone surfaces during the rostral reorientation when defleshing. Denticle width is influenced primarily by tooth size, and correlates well with body size, displaying negative allometry in both groups regardless of taxon or position. When compared, striation widths fall within or below the range of denticle widths extrapolated for similar-sized V. komodoensis individuals. Striation width is directly influenced by the orientation of the carina during feeding, and may underestimate but cannot overestimate denticle width. Although body size can theoretically be estimated solely by a striated tooth mark under ideal circumstances, many caveats should be considered. These include the influence of negative allometry across taxa and throughout ontogeny, the existence of theropods with extreme denticle widths, and the potential for striations to underestimate denticle widths. This method may be useful under specific circumstances, especially for establishing a lower limit body size for potential consumers.

Using striated tooth marks on bone to predict body size in theropod dinosaurs: a model based on feeding observations of Varanus komodoensis, the Komodo monitor

Mesozoic tooth marks on bone surfaces directly link consumers to fossil assemblage formation. Striated tooth marks are believed to form by theropod denticle contact, and attempts have been made to identify theropod consumers by comparing these striations with denticle widths of contemporaneous taxa. The purpose of this study is to test whether ziphodont theropod consumer characteristics can be accurately identified from striated tooth marks on fossil surfaces. We had three major objectives (1) to experimentally produce striated tooth marks and explain how they form; (2) to determine whether body size characteristics are reflected in denticle widths; and (3) to determine whether denticle characters are accurately transcribed onto bone surfaces in the form of striated tooth marks. We conducted controlled feeding trials with the dental analogue Varanus komodoensis (the Komodo monitor). Goat (Capra hircus) carcasses were introduced to captive, isolated individuals. Striated tooth marks were then identified, and striation width, number, and degree of convergence were recorded for each. Denticle widths and tooth/body size characters were taken from photographs and published accounts of both theropod and V. komodoensis skeletal material, and regressions were compared among and between the two groups. Striated marks tend to be regularly striated with a variable degree of branching, and may co-occur with scores. Striation morphology directly reflects contact between the mesial carina and bone surfaces during the rostral reorientation when defleshing. Denticle width is influenced primarily by tooth size, and correlates well with body size, displaying negative allometry in both groups regardless of taxon or position. When compared, striation widths fall within or below the range of denticle widths extrapolated for similar-sized V. komodoensis individuals. Striation width is directly influenced by the orientation of the carina during feeding, and may underestimate but cannot overestimate denticle width. Although body size can theoretically be estimated solely by a striated tooth mark under ideal circumstances, many caveats should be considered. These include the influence of negative allometry across taxa and throughout ontogeny, the existence of theropods with extreme denticle widths, and the potential for striations to underestimate denticle widths. This method may be useful under specific circumstances, especially for establishing a lower limit body size for potential consumers.

The Effects of Biting and Pulling on the Forces Generated during Feeding in the Komodo Dragon (Varanus komodoensis)

In addition to biting, it has been speculated that the forces resulting from pulling on food items may also contribute to feeding success in carnivorous vertebrates. We present an in vivo analysis of both bite and pulling forces in Varanus komodoensis, the Komodo dragon, to determine how they contribute to feeding behavior. Observations of cranial modeling and behavior suggest that V. komodoensis feeds using bite force supplemented by pulling in the caudal/ventrocaudal direction. We tested these observations using force gauges/transducers to measure biting and pulling forces. Maximum bite force correlates with both body mass and total body length, likely due to increased muscle mass. Individuals showed consistent behaviors when biting, including the typical medial-caudal head rotation. Pull force correlates best with total body length, longer limbs and larger postcranial motions. None of these forces correlated well with head dimensions. When pulling, V. komodoensis use neck and limb movements that are associated with increased caudal and ventral oriented force. Measured bite force in Varanus komodoensis is similar to several previous estimations based on 3D models, but is low for its body mass relative to other vertebrates. Pull force, especially in the ventrocaudal direction, would allow individuals to hunt and deflesh with high success without the need of strong jaw adductors. In future studies, pull forces need to be considered for a complete understanding of vertebrate carnivore feeding dynamics.

Cut! Could the dragon do that again, please?

Cut! Could the dragon do that again, please?