Maximum body size among insular Komodo dragon populations covaries with large prey density

This study documents variation in maximum body size of Komodo dragons ( Varanus komodoensis ) among the four extant island populations in Komodo National Park and compares an indirect measure of deer density, the major prey item for large dragons, to differences in maximum body size among islands. The largest 15% of dragons from the large islands of Komodo and Rinca were significantly longer and heavier than the largest 15% of dragons on the small islands of Gili Motang and Nusa Kode. There was a 33% difference in snout vent length (SVL) between dragons found on Komodo and those found on Gili Motang, with mass varying by more than four-fold. Density of deer pellet groups between islands ranged from 5.86±0.75 groups per transect on Gili Motang to 20.73±1.02 groups per transect on Komodo Island. Maximal dragon SVL and mass was highly positively correlated with this index of deer density. Low prey density on the two small islands could constrain body size via energetic constraints. At present we can not deduce if insular body size variation has arisen through genotypic or phenotypic mechanisms.     

Evaluation of three field monitoring-density estimation protocols and their relevance to Komodo dragon conservation

Finding practical ways to robustly estimate abundance or density trends in threatened species is a key facet for effective conservation management. Further identifying less expensive monitoring methods that provide adequate data for robust population density estimates can facilitate increased investment into other conservation initiatives needed for species recovery. Here we evaluated and compared inference-and cost-effectiveness criteria for three field monitoring-density estimation protocols to improve conservation activities for the threatened Komodo dragon (Varanus komodoensis). We undertook line-transect counts, cage trapping and camera monitoring surveys for Komodo dragons at 11 sites within protected areas in Eastern Indonesia to collect data to estimate density using distance sampling methods or the Royle–Nichols abundance induced heterogeneity model. Distance sampling estimates were considered poor due to large confidence intervals, a high coefficient of variation and that false absences were obtained in 45 % of sites where other monitoring methods detected lizards present. The Royle–Nichols model using presence/absence data obtained from cage trapping and camera monitoring produced highly correlated density estimates, obtained similar measures of precision and recorded no false absences in data collation. However because costs associated with camera monitoring were considerably less than cage trapping methods, albeit marginally more expensive than distance sampling, better inference from this method is advocated for ongoing population monitoring of Komodo dragons. Further the cost-savings achieved by adopting this field monitoring method could facilitate increased expenditure on alternative management strategies that could help address current declines in two Komodo dragon populations.     

Blood values in wild and captive Komodo dragons (Varanus komodoensis)

The Komodo dragon (Varanus komodoensis) is the largest living lizard and occupies a range smaller than that of any other large carnivore in the world. Samples from 33 free-ranging animals at five localities in Komodo National Park, Indonesia were evaluated to assess underlying health problems. To build a comparative database, samples from 44 Komodo dragons in both Indonesian and U.S. zoos were also analyzed. Tests performed included complete blood counts, clinical chemistry profiles, vitamin A, D(3), and E analyses, mineral levels, and screening for chlorinated pesticides or other toxins in wild specimens. Blood samples from wild dragons were positive for hemogregarines, whereas captive specimens were all negative. Total white blood cell counts were consistently higher in captive Komodo dragons than in wild specimens. Reference intervals were established for some chemistry analytes, and values obtained from different groups were compared. Vitamin A and E ranges were established. Vitamin D(3) levels were significantly different in Komodo dragons kept in captive, indoor exhibits versus those with daily ultraviolet-B exposure, whether captive or wild specimens. Corrective measures such as ultraviolet-permeable skylights, direct sunlight exposure, and self-ballasted mercury vapor ultraviolet lamps increased vitamin D(3) concentrations in four dragons to levels comparable with wild specimens. Toxicology results were negative except for background-level chlorinated pesticide residues. The results indicate no notable medical, nutritional, or toxic problems in the wild Komodo dragon population. Problems in captive specimens may relate to, and can be corrected by, husbandry measures such as regular ultraviolet-B exposure. Zoo Biol 19:495-509, 2000. Copyright 2000 Wiley-Liss, Inc.     

Ontogenetic differences in the spatial ecology of immature Komodo dragons

The early life-history stages of reptiles are extremely important to an individual's fitness, but in an ecological sense, among the most difficult to observe. Here, we used radio-tracking techniques to describe the differences in movement patterns, habitat use and home range between hatchling and juvenile Komodo dragons Varanus komodoensis on Komodo Island, Indonesia. The movement of hatchlings from their nests was largely linear and suggested a natal dispersal event. The movement patterns of juvenile Komodo dragons exhibited a greater spatial overlap than hatchlings, indicating greater site fidelity and thus use of a more defined activity area. The rates of daily movement were significantly less for hatchlings compared with juvenile dragons. The activity areas of hatchlings were significantly smaller than juvenile dragons. Both age classes preferred utilizing dry monsoon forest compared with other habitat types. Hatchlings were predominantly arboreal compared with juveniles and the degree of arboreal activity was strongly correlated with an individual's size. These distinct differences in spatial ecology between immature life-history stages suggest that different selection pressures may affect different size classes of Komodo dragons.     

Aerobic salivary bacteria in wild and captive Komodo dragons

During the months of November 1996, August 1997, and March 1998, saliva and plasma samples were collected for isolation of aerobic bacteria from 26 wild and 13 captive Komodo dragons (Varanus komodoensis). Twenty-eight Gram-negative and 29 Gram-positive species of bacteria were isolated from the saliva of the 39 Komodo dragons. A greater number of wild than captive dragons were positive for both Gram-negative and Gram-positive bacteria. The average number of bacterial species within the saliva of wild dragons was 46% greater than for captive dragons. While Escherichia coli was the most common bacterium isolated from the saliva of wild dragons, this species was not present in captive dragons. The most common bacteria isolated from the saliva of captive dragons were Staphylococcus capitis and Staphylococcus capitis and Staphylococcus caseolyticus, neither of which were found in wild dragons. High mortality was seen among mice injected with saliva from wild dragons and the only bacterium isolated from the blood of dying mice was Pasteurella multocida. A competitive inhibition enzyme-linked immunosorbent assay revealed the presence of anti-Pasteurella antibody in the plasma of Komodo dragons. Four species of bacteria isolated from dragon saliva showed resistance to one or more of 16 antimicrobics tested. The wide variety of bacteria demonstrated in the saliva of the Komodo dragon in this study, at least one species of which was highly lethal in mice and 54 species of which are known pathogens, support the observation that wounds inflicted by this animal are often associated with sepsis and subsequent bacteremia in prey animals.     

Body temperature and thermoregulation of Komodo dragons in the field

Komodo dragons from hatchlings (≈0.1 kg) to adults (≤80 kg) express the full magnitude of varanid species size distributions. We found that all size groups of dragons regulated a similar preferred body temperature by exploiting a heterogeneous thermal environment within savanna, forest and mangrove habitats. All dragons studied, regardless of size, were able to regulate a daytime active body temperature within the range 34–35.6 °C for 5.1–5.6 h/day. The index of effectiveness of thermoregulation (a numerical rating of thermoregulatory activity) was not different among size groups of dragons. However, the index of closeness of thermoregulation, which rates the variability of body temperature, suggests a greater precision for regulating a preferred body temperature for medium compared to small and large dragons. Reference copper cylinders simulating small, medium and large Komodo dragons heated and cooled at the same rate, whereas actual dragons of all size groups heated faster than they cooled. Larger dragons heated and cooled more slowly than smaller ones. The mean operative environmental temperatures of copper cylinders representing medium sized dragons were 42.5, 32.0 and 29.4° C for savannah, forest and mangrove habitats, respectively. The index for average thermal quality of a habitat as measured by the absolute difference between operative environmental temperature and the dragon’s thermal range suggests the forest habitat offers the highest thermal quality to dragons and the savannah the lowest. The percent of total daytime that the operative environmental temperature was within the central 50% of the body temperatures selected by dragons in a thermal gradient (Phillips, 1984) was 45%, 15%, and 9% for forest, mangrove and savannah, respectively. Forest habitat offers the most suitable thermal environment and provides the greatest number of hours with conditions falling within the dragon’s thermal activity zone.     

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.     

Distribution, seasonal use, and predation of incubation mounds of Orange-footed Scrubfowl on Komodo Island, Indonesia

ABSTRACT.   Megapodes are unique in using only heat from the environment, rather than body heat, to incubate their eggs as well as the precocious independence of their chicks on hatching. Of 22 recognized species of megapodes, 9 are listed as threatened due to factors including habitat loss and fragmentation, and predation on eggs and chicks. Orange-footed Scrubfowl ( Megapodius reinwardt ) are conspicuous components of the Oriental/Austral avifauna that inhabit the monsoon forests of the Lesser Sunda chain of islands in Indonesia. We examined the abundance, patterns of distribution, physical characteristics, seasonal activity, and predation risk of incubation mounds of Orange-footed Scrubfowl on Komodo Island in eastern Indonesia. We surveyed 13 valleys on Komodo Island from April 2002 to January 2005 and located 113 tended and 107 untended incubation mounds. Densities of scrubfowl mounds in our study were similar to that reported by investigators during the 1970s, suggesting little change in the scrubfowl population since then. Most scrubfowl mounds were on sandy or loamy soils in open monsoon forest with little overhead shade, and placement of mounds in such areas may ensure adequate temperatures for egg incubation. Although some mounds were tended during all months, mound use peaked during the late wet season in March. During the dry season (April–November), only a few mounds were tended. Komodo dragons ( Varanus komodoensis ) and wild pigs ( Sus scrofa ) were the primary predators of scrubfowl eggs, with no indication of egg predation by humans. The valley with the largest number of untended mounds in our study also had the largest number of active Komodo dragon nests. This suggests an effect of Komodo dragons on scrubfowl numbers, but additional study is needed.     

Evaluating environmental,demographic and genetic effects on population‐level survival in an island endemic

The population dynamics of island species are considered particularly sensitive to variation in environmental, demographic and/or genetic processes. However, few studies have attempted to evaluate the relative importance of these processes for key vital rates in island endemics. We integrated the results of long‐term capture–mark–recapture analysis, prey surveys, habitat quality assessments and molecular analysis to determine the causes of variation in the survival rates of Komodo dragons Varanus komodoensis at 10 sites on four islands in Komodo National Park, Indonesia. Using open population capture–mark–recapture methods, we ranked competing models that considered environmental, ecological, genetic and demographic effects on site‐specific Komodo dragon survival rates. Site‐specific survival rates ranged from 0.49 (95% CI: 0.33–0.68) to 0.92 (0.79–0.97) in the 10 study sites. The three highest‐ranked models (i.e. ΔQAIC_c < 2) explained ～70% of variation in Komodo dragon survival rates and identified interactions between inbreeding coefficients, prey biomass density and habitat quality as important explanatory variables. There was evidence of additive effects from ecological and genetic (e.g. inbreeding) processes affecting Komodo dragon survival rates. Our results indicate that maintaining high ungulate prey biomass and habitat quality would enhance the persistence of Komodo dragon populations. Assisted gene flow may also increase the genetic and demographic viability of the smaller Komodo dragon populations.     

Genetic divergence and units for conservation in the Komodo dragon Varanus komodoensis

In the past decade much attention has focused on the role that genetics can play in the formation of management strategies in conservation. Here, we describe genetic diversity in the world''s largest lizard, the Komodo dragon (Varanus komodoensis), examining the evolutionary relationships and population genetic history of the four islands in south-east Indonesia, which form the vast majority of its range. We identify distinct genetic groups for conservation. The population on the island of Komodo shows by far the largest values of genetic divergence and is proposed that it should be a separate conservation management unit. Other populations, surviving either on small islands with substantially reduced genetic variability, or in isolated patches, are identified as particularly vulnerable to stochastic threats and habitat loss. Our results provide an example of how data defining intraspecific levels of genetic divergence can provide information to help management plans, ensure the maintenance of genetic variability across populations and identify evolutionary potential within endangered species.     

Nutritional Consequences of Experimentally Introduced Tourism in Brown Bears

Abstract: Although numerous studies have documented behavioral effects of nature-based tourism on wildlife populations, few studies have determined whether behavioral changes translate to effects on individual condition and population health. This issue is currently a concern for wildlife managers in Alaska, USA, and Canada where bear viewing is a rapidly growing industry expanding into previously undisturbed bear habitats. Rather than record observations at long established tourism sites, we experimentally introduced bear viewing into 2 relatively undisturbed brown bear (Ursus arctos) populations in south-central Alaska. We examined the nutritional consequences of behavioral changes induced by the presence and activity of bear viewers for bears feeding on early summer vegetation and late-summer salmon (Oncorhynchus kisutch and O. nerka). We used Global Positioning System collars, monitored food resource availability, and quantified individual resource use and condition for a year prior to and during the introduction of bear viewing. Though bear viewing altered spatiotemporal resource use in all treatments, total resource use declined only when we exposed bears to 24-hour daily human activity. Energy expenditure, indexed as daily travel distances, was significantly higher when bears responded by altering spatial rather than temporal resource use. However, body weight and composition were unaffected by all treatments as bears shifted their foraging to other locations or times. Managers can minimize nutritional impacts of bear-viewing programs by avoiding spatial displacement and providing predictable time periods when bears can access food resources free of human activity. Bears in this study exhibited a high degree of behavioral plasticity, which may be an important factor in identifying flagship species for sustainable ecotourism programs.     

Genetic structure and gene flow among Komodo dragon populations inferredby microsatellite loci analysis

A general concern for the conservation of endangered species is the maintenance of genetic variation within populations, particularly when they become isolated and reduced in size. Estimates of gene flow and effective population size are therefore important for any conservation initiative directed to the long-term persistence of a species in its natural habitat. In the present study, 10 microsatellite loci were used to assess the level of genetic variability among populations of the Komodo dragon Varanus komodoensis. Effective population size was calculated and gene flow estimates were compared with palaeogeographic data in order to assess the degree of vulnerability of four island populations. Rinca and Flores, currently separated by an isthmus of about 200 m, retained a high level of genetic diversity and showed a high degree of genetic similarity, with gene flow values close to one migrant per generation. The island of Komodo showed by far the highest levels of genetic divergence, and its allelic distinctiveness was considered of great importance in the maintenance of genetic variability within the species. A lack of distinct alleles and low levels of gene flow and genetic variability were found for the small population of Gili Motang island, which was identified as vulnerable to stochastic threats. Our results are potentially important for both the short- and long-term management of the Komodo dragon, and are critical in view of future re-introduction or augmentation in areas where the species is now extinct or depleted.     

Development of a multiplex PCR assay for fine-scale population genetic analysis of the Komodo monitor Varanus komodoensis based on 18 polymorphic microsatellite loci

Multiplex PCR assays for the coamplification of microsatellite loci allow rapid and cost-effective genetic analyses and the production of efficient screening protocols for international breeding programs. We constructed a partial genomic library enriched for di-nucleotide repeats and characterized 14 new microsatellite loci for the Komodo monitor (or Komodo dragon, Varanus komodoensis). Using these novel microsatellites and four previously described loci, we developed multiplex PCR assays that may be loaded on a genetic analyser in three separate panels. We tested the novel set of microsatellites for polymorphism using 69 individuals from three island populations and evaluated the resolving power of the entire panel of 18 loci by conducting (i) a preliminary assignment test to determine population(s) of origin and (ii) a parentage analysis for 43 captive Komodo monitors. This panel of polymorphic loci proved useful for both purposes and thus can be exploited for fine-scale population genetic analyses and as part of international captive breeding programs directed at maintaining genetically viable ex situ populations and reintroductions.     

Deathly Drool: Evolutionary and Ecological Basis of Septic Bacteria in Komodo Dragon Mouths

Komodo dragons, the world''s largest lizard, dispatch their large ungulate prey by biting and tearing flesh. If a prey escapes, oral bacteria inoculated into the wound reputedly induce a sepsis that augments later prey capture by the same or other lizards. However, the ecological and evolutionary basis of sepsis in Komodo prey acquisition is controversial. Two models have been proposed. The “bacteria as venom” model postulates that the oral flora directly benefits the lizard in prey capture irrespective of any benefit to the bacteria. The “passive acquisition” model is that the oral flora of lizards reflects the bacteria found in carrion and sick prey, with no relevance to the ability to induce sepsis in subsequent prey. A third model is proposed and analyzed here, the “lizard-lizard epidemic” model. In this model, bacteria are spread indirectly from one lizard mouth to another. Prey escaping an initial attack act as vectors in infecting new lizards. This model requires specific life history characteristics and ways to refute the model based on these characteristics are proposed and tested. Dragon life histories (some details of which are reported here) prove remarkably consistent with the model, especially that multiple, unrelated lizards feed communally on large carcasses and that escaping, wounded prey are ultimately fed on by other lizards. The identities and evolutionary histories of bacteria in the oral flora may yield the most useful additional insights for further testing the epidemic model and can now be obtained with new technologies.     

Evolutionary and ecological traps for brown bears Ursus arctos in human‐modified landscapes

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Variation in behavioral traits of two frugivorous mammals may lead to differential responses to human disturbance

Human activities can lead to a shift in wildlife species’ spatial distribution. Understanding the specific effects of human activities on ranging behavior can improve conservation management of wildlife populations in human‐dominated landscapes. This study evaluated the effects of forest use by humans on the spatial distribution of mammal species with different behavioral adaptations, using sympatric western lowland gorilla and central chimpanzee as focal species. We collected data on great ape nest locations, ecological and physical variables (habitat distribution, permanent rivers, and topographic data), and anthropogenic variables (distance to trails, villages, and a permanent research site). Here, we show that anthropogenic variables are important predictors of the distribution of wild animals. In the resource model, the distribution of gorilla nests was predicted by nesting habitat distribution, while chimpanzee nests were predicted first by elevation followed by nesting habitat distribution. In the anthropogenic model, the major predictors of gorilla nesting changed to human features, while the major predictors of chimpanzee nesting remained elevation and the availability of their preferred nesting habitats. Animal behavioral traits (body size, terrestrial/arboreal, level of specialization/generalization, and competitive inferiority/superiority) may influence the response of mammals to human activities. Our results suggest that chimpanzees may survive in human‐encroached areas whenever the availability of their nesting habitat and preferred fruits can support their population, while a certain level of human activities may threaten gorillas. Consequently, the survival of gorillas in human‐dominated landscapes is more at risk than that of chimpanzees. Replicating our research in other sites should permit a systematic evaluation of the influence of human activity on the distribution of mammal populations. As wild animals are increasingly exposed to human disturbance, understanding the resulting consequences of shifting species distributions due to human disturbance on animal population abundance and their long‐term survival will be of growing conservation importance.     

Socio-economics and wildlife conservation of a peri-urban national park in central Kenya

The support, perceptions, values and aspiration of the local people, especially those who neighbour protected areas is increasingly recognised as critical for long-term biodiversity conservation in many countries. As such, conservation agencies are keen to understand the linkages and interactions between wildlife conservation and socio-economics of local communities. We assessed the socio-economic, demographic factors and livelihood status of the households of communities bordering Oldoinyo Sapuk National Park in central Kenya, and evaluated their willingness to support wildlife conservation. We randomly interviewed households that neighbour the Park on a wide range of socio-economic, livelihood and demographic factors. The majority of respondents were small-scale farmers practicing crop farming and livestock keeping. Most respondents considered the Park to have great potential for the future economic development of the area, especially through ecotourism as a potential avenue to support local livelihoods.     

Predation of incubator birds (Megapodius freycinet) by Komodo dragons (Varanus komodoensis)

A short study was made of the nesting habits of a species of megapode, Megapodius freycinet on the island of Komodo. The birds were found to build giant nesting mounds (0.95 × 7.15 m) in scrub and woodland close to the seashore or alongside river beds; 23 active and 19 abandoned mounds were located in an area of 250 ha.
The mounds were apparently added to from year to year, and consisted of dry sand and soil, plus leaves and sticks in most cases. The upper parts of the mounds were invariably exposed to the direct rays of the sun; the temperature within the active mounds was 33–37° C and slightly less in mounds not in use by the birds. According to the local people the nesting season of the megapodes extended from late August to November, and the chicks were hatched before the arrival of the rains in December. Up to four eggs had been recovered from a single mound.
The active mounds were all spaced apart by at least a 100 m, and the closest ones were usually separated by a river bed. This pattern suggested the species was territorial. One or two birds were often seen scraping at a mound or feeding close by, and their behaviour was recorded.
The Komodo dragon, Varanus komodoensis , was a frequent visitor to the nesting mounds of the megapodes, and was able to burrow into them and steal the eggs.     

Approaches to measuring the effects of human disturbance on birds

Human recreational activities are often considered as potential threats to biodiversity, by restricting animals' access to resources that otherwise would be exploited. Because access to wildlife areas is one major means of increasing their public value, and hence the pressure to conserve them, it is clearly critical to be able to identify accurately when human presence is a threat to conservation and when it is not. A wide range of methods have been used to assess the impacts of human disturbance on wildlife and these methods are summarized here. The type of method used depends principally on whether the disturbance issue relates to a particular site, a particular group of individuals or whole populations. Within these categories, both comparative and experimental approaches have been used to assess behavioural, distributional, demographic and population responses to human presence. Examples of each approach are given here, together with an assessment of the information each method provides.     

Predicting the gender of subadult Komodo dragons (Varanus komodoensis) using two-dimensional ultrasound imaging and plasma testosterone concentration

Six yearling Komodo dragons (Varanus komodoensis) underwent ultrasound examinations every three months over a nine-month period to assess the value of two-dimensional ultrasound imaging technology for the purpose of determining gender in this highly monomorphic species. Beginning at an age of 28 months, ovarian structures could be imaged, allowing a prediction of gender based on the presence or absence of ovarian follicles. Simultaneously, plasma testosterone concentration was measured monthly on each individual during the study period. Beginning at an age of 24 months, putative males had consistently elevated plasma testosterone values that were ∼40–60 times greater than those of putative females. In a blind comparison, predictions of gender based on plasma testosterone values matched the prediction of sex based on ultrasound imaging for all six individuals. The results indicate that measurements of circulating testosterone and two-dimensional ultrasound imaging together appear useful in the noninvasive determination of gender in juvenile V. komodoensis at just over two years of age. © 1996 Wiley-Liss, Inc.