Article

Near fixation of melanism in leopards of the Malay Peninsula

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Abstract

Melanistic leopards Panthera pardus are common in south-east Asian forests but the exact frequency of this variant phenotype is difficult to assess. Records from camera-trapping studies conducted at 22 locations in Peninsular Malaysia and southern Thailand between 1996 and 2009 show that only melanistic leopards were present in samples south of the Isthmus of Kra. During 42 565 trap-nights, we collected 445 photos of melanistic leopards and 29 photos of the spotted or non- melanistic morph. All 29 photos of spotted leopards came from study sites north of the Isthmus. These results indicate that this recessive trait may be nearly fixed in P. pardus populations of the Malay Peninsula, suggesting a unique evolutionary history of leopards in the region. Assuming a very small effective population size (Ne = 100) and a high initial allelic frequency, at least 1000 years would be expected to elapse until a neutral allele became fixed. The severe bottleneck implied by this scenario provides a testable hypothesis that can be addressed using molecular markers and evidence of past glacioeustatic changes across the region. Although natural selection might lead to rapid fixation of melanism within Malayan leopards, had their effective population size been much larger (e.g. Ne = 5000) and stable, with a lower allelic frequency, the fixation would require a longer time span (e.g. 20 000 years) if induced by genetic drift alone.

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... In the cat family (Felidae), melanism is quite common, having been reported in 13 of 37 extant species (Table 1). Although such darkened pelage reaches considerably high frequencies in some cat species [4], supporting the notion that this phenotype may be adaptive in some contexts, still little is known about its evolutionary history and ecological/behavioral significance in any felid. Initial molecular analyses have revealed that melanism arose multiple times in the Felidae, with three different mutations being implicated in this phenotype in distinct species [5]. ...
... In contrast, ASIP-induced melanism can reach very high frequency in some felid populations, suggesting that this trait may be adaptive or at least neutral. Such a pattern is particularly noticeable in the case of leopards from the Malay Peninsula, where melanism approaches fixation [4]. Using samples from this very region (seeTable 2), we show here that ASIP is implicated in this mutant phenotype. ...
... Using samples from this very region (seeTable 2), we show here that ASIP is implicated in this mutant phenotype. Although we have shown that this near fixation may have been caused by genetic drift over a long period of time [4], this would be very unlikely if the trait was deleterious. Moreover, such high frequency would be much more quickly achieved if the trait was favorable, and therefore driven to near fixation by natural selection. ...
Article
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The occurrence of melanism (darkening of the background coloration) is documented in 13 felid species, in some cases reaching high frequencies at the population level. Recent analyses have indicated that it arose multiple times in the Felidae, with three different species exhibiting unique mutations associated with this trait. The causative mutations in the remaining species have so far not been identified, precluding a broader assessment of the evolutionary dynamics of melanism in the Felidae. Among these, the leopard (Panthera pardus) is a particularly important target for research, given the iconic status of the 'black panther' and the extremely high frequency of melanism observed in some Asian populations. Another felid species from the same region, the Asian golden cat (Pardofelis temminckii), also exhibits frequent records of melanism in some areas. We have sequenced the coding region of the Agouti Signaling Protein (ASIP) gene in multiple leopard and Asian golden cat individuals, and identified distinct mutations strongly associated with melanism in each of them. The single nucleotide polymorphism (SNP) detected among the P. pardus individuals was caused by a nonsense mutation predicted to completely ablate ASIP function. A different SNP was identified in P. temminckii, causing a predicted amino acid change that should also induce loss of function. Our results reveal two additional cases of species-specific mutations implicated in melanism in the Felidae, and indicate that ASIP mutations may play an important role in naturally-occurring coloration polymorphism.
... The need for white markings as a method of visual communication in low visibility conditions may limit the melanistic phenotype in most felid species (see [1,3]). However, if species or populations are diurnal, such as jaguarundi [49] and leopards [50,51,52], respectively, then melanism may persist with limited negative consequences on communication. ...
... In leopards the frequency of melanistic individuals varies significantly among populations. A very low incidence of melanism has been observed in the open savannas of Africa (1.25%) [1], but reaches 50% of frequency to near fixation in moist forests of the Malay Peninsula [1,4,52]. Interestingly, while populations found in the Malay Peninsula Forests are mainly diurnal [51], African populations display diurnal and nocturnal activity [53]. ...
... However, if natural selection provided adaptive advantages to some polymorphisms [56], anthropogenic changes to the environment may contribute to change the circadian activity patterns of the populations [57], in turn affecting the frequency of melanistic and non-melanistic individuals. This might lead to the disappearance of the "normal" phenotype, similar to the case of leopards in Southeast Asia [52], or the disappearance of melanistic individuals, as in the case of Southern tigrina in southern Brazil ( [23,55], Unpublished data). Therefore, studies of circadian activity that include changes in prey availability and the in the abundance of other carnivore species in a community may contribute to better understand the observed variation in melanism frequencies in felid populations, and ultimately to the conservation of these threatened taxa [58]. ...
Article
Melanism in the cat family has been associated with functions including camouflage, ther-moregulation and parasite resistance. Here we investigate a new hypothesis proposing that the evolution of melanism in cats has additionally been influenced by communication functions of body markings. To evaluate this hypothesis, we assembled a species-level data set of morphological (body marks: white marks on the backs of ears) and ecological (circadian activity: arrhythmic/nocturnal, and environmental preference: open/closed) characteristics that could be associated with communication via body markings, and combined these data with a dated molecular phylogeny. Next, we tested the association between melanism and communication, first by relating species' body marks with their ecological conditions, using a Bayesian implementation of the threshold model. Second, to explore the evolution of characteristics potentially influencing melanism in cat species, we modeled their evolution relative to melanism using models of coordinated vs. independent character changes. Our results suggest that white marks are associated with intraspecific communication between individuals that have non-melanistic phenotypes, as well as towards melanistic individuals (without white marks). The absence of white marks in a melanistic individual tends to be a limiting condition for intraspecific visual communication at night, resulting in an evolutionary dilemma for these species, i.e. to be almost invisible at night, but not to communicate visually. The comparative analysis of several evolutionary models indicated more support for the evolution of melanism being coordinated with the evolution of arrhythmic activity and white marks on the backs of ears.
... Melanism in leopards appears to be most pronounced in one particular region of its distributional range, south of the Isthmus of Kra on mainland Southeast Asia encompassing Peninsular Malaysia and Southern Thailand. To date, almost all observations of leopards in this region have been of melanistic individuals (Kawanishi et al. 2010). Not only is this a unique phenomenon for leopards, it is also possibly the only place in the world where an entire animal population is almost completely composed of the melanistic form of a species. ...
... Not only is this a unique phenomenon for leopards, it is also possibly the only place in the world where an entire animal population is almost completely composed of the melanistic form of a species. This peculiar circumstance may be the result of 1) a founder effect following a historical population crash, 2) natural selection driven by competition with tigers (Panthera tigris) and low light conditions caused by a dense forest canopy, or 3) random genetic drift (Kawanishi et al. 2010). The only other known populations where melanistic leopards may constitute a large proportion can be found in Java and central Thailand; however, in these areas, spotted leopards have been recorded at a greater frequency than in Malaysia (Pocock 1930;Gippoliti and Meijaard 2007;Steinmetz et al. 2007Steinmetz et al. , 2009. ...
... However, this approach may be unsuitable for large carnivores because of non-random movement of individuals (Foster and Harmsen 2012). Nevertheless, although leopards in Malaysia often appear uniformly black in camera trap photos, it is possible to see the more heavily pigmented rosette markings within their pelage under certain lighting conditions (Kawanishi et al. 2010, Schneider et al. 2012. The rosette markings are especially visible when viewed on images where an infrared flash has been used to illuminate the animal (Fig. 1). ...
Article
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To date, leopards (Panthera pardus) in Peninsular Malaysia have been overlooked by large carnivore researchers. This is in part due to the country's unique population of individuals that are almost all melanistic, which makes it nearly impossible to identify individuals using camera traps for estimating leopard density. We discovered a novel modification to infrared flash camera traps, which forces the camera into night mode, that allows us to consistently and clearly see the spots of a melanistic leopard. The aim of this project was 1) to determine the feasibility of identifying melanistic leopards with confidence using infrared flash camera traps, and 2) to establish a density estimate for the leopard population in a wildlife corridor in Malaysia using maximum likelihood and Bayesian spatially explicit capture-recapture (SECR) models. Both SECR approaches yielded a leopard density of approximately 3 individuals/100 km2. Our estimates represent the first density estimate of leopards in Malaysia and arguably, the world's first successful attempt to estimate the population size of a species with melanistic phenotypes. Because we have demonstrated that melanistic leopards can be monitored with confidence using infrared cameras, future studies should employ our approach instead of relying on scars or body shape for identification. Ultimately, our approach can facilitate more accurate assessments of leopard population trends, particularly in regions where melanistic phenotypes largely occur. © 2015 The Wildlife Society.
... The black leopard is common in the equatorial rainforest of Malaya, in the forests and mountains of Java, and in the tropical rainforest on the slopes of some African mountains, such as Mount Kenya [3,4]. They have also been reported to occur in China, Myanmar, Assam, Thailand, and Nepal [5]. Interestingly, a study in 2009 reported that leopards south of the Isthmus of Kra on mainland Southeast Asia (Peninsular Malaysia and Southern Thailand) are all melanistic. ...
... The spotted leopard is rare in Peninsular Malaysia -so far it has been reported only in two other written works: one in Endau Rompin National Park, in the southern state of Johor [8]; and the other mentioned anecdotally (in an old identification book) as being outnumbered by 'conventionally colored leopards' in Malaya [9]. This is despite the extensive camera trapping efforts in Peninsular Malaysia, all of which (with the exception of the above two studies) have recorded the presence of only the melanistic form [6,[10][11][12][13][14][15]. A study in 2010 examined the frequency of leopard melanism in 22 locations in Peninsular Malaysia and southern Thailand [5] (Fig. 2). Out of 42,565 trap-nights, 445 photos of melanistic leopards and 29 photos of the spotted morph were collected; all photos of the spotted leopard were taken from study areas north of the Isthmus of Kra, which is the narrow neck of southern Myanmar and Thailand and a well-known zoogeographical boundary [16]. ...
... All these suggest that spotted leopards are at the very least rare in Peninsular Malaysia. Kawanishi (2010) suggested that the black morph has become genetically fixed south of the Isthmus of Kra due to an historical bottleneck event. Subsequently, genetic drift or natural selection or both could explain the fixation of melanism in leopards in Peninsular Malaysia. ...
Article
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We discovered the presence of two individual spotted leopards Panthera pardus in Ulu Muda Forest Reserve, a previously under-researched selectively logged rainforest of Peninsular Malaysia. These findings are unexpected, because only two other studies have detected the spotted morph amongst many other melanistic leopards caught on camera traps in Peninsular Malaysia. We discuss the implications of our findings with respect to the prevalence of melanism among leopards in the region. © Cedric Tan Kai Wei, Jonathan Moore, Salman bin Saaban, Ahimsa Campos-Arceiz and David W. Macdonald.
... These patterns are thought to correspond with differing vegetation types and light levels in order to conceal the animal from prey and possibly other predators (Allen et al. 2010;Kingdon et al. 2013). This adaptive explanation is supported by the frequent occurrence of melanistic leopards in humid habitats such as the Malayan peninsula (Kawanishi et al. 2010;Schneider et al. 2012). The frequency of 'black panthers' dramatically decreases across more arid regions (Kawanishi et al. 2010). ...
... This adaptive explanation is supported by the frequent occurrence of melanistic leopards in humid habitats such as the Malayan peninsula (Kawanishi et al. 2010;Schneider et al. 2012). The frequency of 'black panthers' dramatically decreases across more arid regions (Kawanishi et al. 2010). The release of eumelanin (black pigmentation) into mammalian pelage is known to be regulated by the extension gene and phaeomelanin (yellow-red pigmentation) by the agouti gene (Fontanesi et al. 2009). ...
... Such inbreeding effects are thought to result in the presence of the very rare king cheetah (Acinonyx jubatus) and white lion colour morphs in southern Africa (Sunquist & Sunquist 2014); it has also been documented with leopards in the Malaysian peninsula, where the rapid near fixation of melanism occurred following population fragmentation (Hedges et al. 2015;Kawanishi et al. 2010). Similarly, Haag et al. (2010) reported genetic drift within small fragmented jaguar populations over a relatively short time frame and McManus et al. (2015) showed that leopard populations can become isolated within a few generations. ...
Article
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Background: Leopards (Panthera pardus) show genetically determined colour variation. Erythristic (strawberry) morphs, where individuals are paler and black pigment in the coat is replaced by a red-brown colour, are exceptionally rare in the wild. Historically, few records exist, with only five putative records known from India.Objectives: To record the presence of erythristic leopards in our study site (Thaba Tholo Wilderness Reserve, Mpumalanga) and to collate records from across South Africa. Method: A network of camera traps was used to record individual leopards at Thaba Tholo. We also surveyed local experts, searched the popular South African press, and used social media to request observations.Results: Two out of 28 individual leopards (7.1%) recorded in our study site over 3 years were of this colour morph. We obtained records of five other erythristic leopards in the North West and Mpumalanga regions, with no reports outside of this population.Conclusions: Erythristic leopards are widely dispersed across north-east South Africa, predominantly in the Lydenburg region, Mpumalanga. The presence of this rare colour morph may reflect the consequences of population fragmentation.
... The occurrence of melanism is rather common in Felidae, having been documented in 13 of the 38 felid species, evolved independently at least eight times within the family [11][12][13], in some cases reaching very high frequencies in natural populations [14,15]. In none of them has it reached fixation, but rather always exists as a polymorphic phenotype, and it is present only in two species of Panthera: the leopard (P. ...
... In fact, this study has provided a characterization of the spatial distribution of melanism in leopards and jaguars and demonstrated that such distribution is nonrandom. Recent analyses have indicated that melanism can reach very high frequencies in some leopard populations (e.g., Southeast Asia reported by [14]). In addition, there have been confirmed reports of melanistic leopards in India, Abyssinia, and Ethiopian Highlands, Java and Malaysia, Aberdare Mount Kenya, Highlands of Nepal, as well as a doubtful occurrence in South Africa [19,26,28,54]. ...
... When the distribution of melanism in leopards is examined more closely, Southeast Asia emerges as a particularly interesting region. Our data support the findings reported by [14] and [15], showing that melanism is almost fixed in areas south of the Isthmus of Kra (Thailand/Malaysia). This study obtained only two records of nonmelanistic animals south of the Isthmus, while in more northerly areas both phenotypes appear at similar frequencies. ...
... Adding to this is the difficulty of conducting field studies in thick tropical rainforest compared to other leopard habitats and lack of funding or interest compared to more charismatic species like the tiger. Interestingly an overwhelming majority of individuals documented in the peninsula are of the black or melanistic variety (Kawanishi et al. 2010). It is this characteristic of Malaysian leopards that this paper shall focus on, discussing the various theories explaining this unusual phenomenon and whether it is a sufficiently unique attribute to make the black panthers here evolutionary distinct from other Southeast Asian leopards. ...
... This makes the concentration of black leopards in Malaysia a phenomenon that warrants further investigation. This topic was extensively discussed and the subject of Kawanishi et al. (2010), which collated the results of past surveys in the country evidencing the overwhelming prevalence of black leopards in the peninsula. Here I summarize the theories and elaborate on those I believe to be most likely explanations. ...
... The unique history and situation of Peninsular Malaysia in Sundaland has already been discussed, and the existence of a bottleneck at the Kra Isthmus mentioned as a potential inhibitor of genetic exchange for leopards with the rest of mainland Southeast Asia. Kawanishi et al. (2010) conjectured that an isolated small population would have made it possible for genetic drift alone to account for melanism becoming fixed over a period of 20,000-40,000 years. ...
Article
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Modern leopards originated in Africa less than a million years ago and dispersed east through Central Asia, the Indian subcontinent up to the Far East and Southeast Asia, where it has a discontinuous distribution today. Scarce resources, intraguild competition and the eruption of Mt Toba may have led to extinction of leopards on Sumatra. In Peninsular Malaysia the range and population of leopards have decreased since the last century along with reduction in suitable forest habitat, to an estimated 500-700 individuals today. The fact that nearly all leopards on the peninsula are melanistic is an unusual phenomenon attributed to genetic drift and natural selection, the latter likely driven by advantage in concealment from tigers in a thickly forested environment. Melanism alone does not make leopards in Malaysia genetically distinct from other Indochinese leopards (Panthera pardus delacouri) to warrant classification as a different subspecies, but it does make them a globally unique subpopulation worthy of conservation and protection.
... The need for white markings as a method of visual communication in low visibility conditions may limit the melanistic phenotype in most felid species (see [1,3]). However, if species or populations are diurnal, such as jaguarundi [49] and leopards [50,51,52], respectively, then melanism may persist with limited negative consequences on communication. ...
... In leopards the frequency of melanistic individuals varies significantly among populations. A very low incidence of melanism has been observed in the open savannas of Africa (1.25%) [1], but reaches 50% of frequency to near fixation in moist forests of the Malay Peninsula [1,4,52]. Interestingly, while populations found in the Malay Peninsula Forests are mainly diurnal [51], African populations display diurnal and nocturnal activity [53]. ...
... However, if natural selection provided adaptive advantages to some polymorphisms [56], anthropogenic changes to the environment may contribute to change the circadian activity patterns of the populations [57], in turn affecting the frequency of melanistic and non-melanistic individuals. This might lead to the disappearance of the "normal" phenotype, similar to the case of leopards in Southeast Asia [52], or the disappearance of melanistic individuals, as in the case of Southern tigrina in southern Brazil ( [23,55], Unpublished data). Therefore, studies of circadian activity that include changes in prey availability and the in the abundance of other carnivore species in a community may contribute to better understand the observed variation in melanism frequencies in felid populations, and ultimately to the conservation of these threatened taxa [58]. ...
Article
Full-text available
Melanism in the cat family has been associated with functions including camouflage, ther-moregulation and parasite resistance. Here we investigate a new hypothesis proposing that the evolution of melanism in cats has additionally been influenced by communication functions of body markings. To evaluate this hypothesis, we assembled a species-level data set of morphological (body marks: white marks on the backs of ears) and ecological (circadian activity: arrhythmic/nocturnal, and environmental preference: open/closed) characteristics that could be associated with communication via body markings, and combined these data with a dated molecular phylogeny. Next, we tested the association between melanism and communication, first by relating species' body marks with their ecological conditions, using a Bayesian implementation of the threshold model. Second, to explore the evolution of characteristics potentially influencing melanism in cat species, we modeled their evolution relative to melanism using models of coordinated vs. independent character changes. Our results suggest that white marks are associated with intraspecific communication between individuals that have non-melanistic phenotypes, as well as towards melanistic individuals (without white marks). The absence of white marks in a melanistic individual tends to be a limiting condition for intraspecific visual communication at night, resulting in an evolutionary dilemma for these species, i.e. to be almost invisible at night, but not to communicate visually. The comparative analysis of several evolutionary models indicated more support for the evolution of melanism being coordinated with the evolution of arrhythmic activity and white marks on the backs of ears.
... Recent analyses have indicated that melanism can reach very high frequencies in some leopard populations (e.g. Southeast Asia [17][18]). In addition, there have been confirmed reports of melanistic leopards in India [15; 19-20], Ethiopia [15; 21], Java and Malaysia [17-18; 22-25], Aberdare Mountains in Kenya [15] and Nepal [26], as well as a potential occurrence in South Africa [15; 27]. ...
... We explored this comparison in more depth using the Southeast Asia data set (S5 Figpanel E). Our data support the observations reported by [17] and [18], showing that leopard melanism is almost fixed in areas south of the Isthmus of Kra (Thailand/Malaysia). We obtained only two records of non-melanistic animals south of the Isthmus (the same was reported by [59]), while in more northerly areas both phenotypes appear at similar frequencies. ...
Article
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The geographic distribution and habitat association of most mammalian polymorphic phenotypes are still poorly known, hampering assessments of their adaptive significance. Even in the case of the black panther, an iconic melanistic variant of the leopard (Panthera pardus), no map exists describing its distribution. We constructed a large database of verified records sampled across the species' range, and used it to map the geographic occurrence of mela-nism. We then estimated the potential distribution of melanistic and non-melanistic leopards using niche-modeling algorithms. The overall frequency of melanism was ca. 11%, with a significantly non-random spatial distribution. Distinct habitat types presented significantly different frequencies of melanism, which increased in Asian moist forests and approached zero across most open/dry biomes. Niche modeling indicated that the potential distributions of the two phenotypes were distinct, with significant differences in habitat suitability and rejection of niche equivalency between them. We conclude that melanism in leopards is strongly affected by natural selection, likely driven by efficacy of camouflage and/or thermo-regulation in different habitats, along with an effect of moisture that goes beyond its influence on vegetation type. Our results support classical hypotheses of adaptive coloration in animals (e.g. Gloger's rule), and open up new avenues for in-depth evolutionary analyses of melanism in mammals.
... Melanism has �een defined as visi�le darker coat colouration due to higher concentration of the melanin pigments compared to the representative form of an individual within a species or population (Kettlewell 1973, Ma� jerus 1998, Kawanishi et al. 2010. Melanism in coat colouration is a quite common genetic polymorphism in wild felids (Eizirik et al. 2003, Schneider et al. 2012. ...
... Melanism in coat colouration is a quite common genetic polymorphism in wild felids (Eizirik et al. 2003, Schneider et al. 2012. Presence of �oth large and small wild cat species with melanistic coat pattern were reported from South�east Asia and the Indian su�continent (Kawanishi et al. 2010, Bashir et al. 2011, Das et al. 2012. The jungle cat is the largest mem�er of the genus Felis, which consists of small cats in the family Felidae (Wozencraft 2005, John� son et al. 2006. ...
Article
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The jungle cat Felis chaus is a habitat generalist and a quite commonly found wild cat on the Indian subcontinent. Earlier studies from South-east Asia and the Indian subcontinent reported melanism in wild felids such as in leopards Panthera pardus, Asiatic golden cats Catopuma temminckii, leopard cats Prionailurus bengalensis etc. A recently conducted camera trapping exercise confirmed the presence of a few melanistic jungle cats from the semi-arid dry deciduous forests of Ranthambhore Tiger Reserve RTR in the state of Rajasthan, north-western India. This is the first photographic report of the melanistic form of the jungle cat from this part of the world.
... The hair, skin and eye color are primarily determined by the quantity and distribution of eumelanin (black and brown coloration) and phaeomelanin (red and yellow coloration), which are produced by melanocytes present in the epithelium, iris, and hair (Prota 1980;Sanchez-Ferrer et al. 1995). The occurrence of phenotypic variations in the coat of animals is related to geographic distribution, seasonal climatic variations, age, sex, weight, and other factors (Ortolani 1999;Stoner et al. 2003;Kawanishi et al. 2010;Ancilloto & Mori 2017). Anomalous colors occur when integumentary pigments are present in excess or in deficient amounts in parts or in totality of the body; white is the complete lack of pigment (Prota 1980;Ortolani 1999;Caro 2005;Fertl & Rosel 2002). ...
... Considering the number of records reported the analysis of the patterns of genetic diversity within and among local populations would be important. The occurrence of the leucistic phenotype observed in the Oriximiná and Caraça populations may represent an opportunity to investigate the possibility of establishing a recessive trait in these populations, such as the near fixation of melanism, another recessive color anomaly, observed in leopards of the Malay Peninsula (Kawanishi et al. 2010). On the other hand, the occurrence of different coloration phenotypes in tayras throughout its distribution area, provides an opportunity to investigate possible adaptive advantages of these phenotypes (Silva et al. 2016) in relation to the ecological conditions present in their areas of occurrence. ...
Article
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The occurrence of anomalous coloration (albinism, leucism and melanism) in mammals is a rare phenomenon in nature, but this phenomenon has been reported for several species of mammals. In this study, we report on the occurrence of leucism in Eira barbara by examining three road-killed individuals and two sightings of live animals in Reserva Particular do Patrimônio Natural Santuário do Caraça, southeastern Brazil. In addition, we examined tayra specimens housed in mammal collections from Brazil and USA. The animals found dead and those sighted had a whitish yellow fur on the body and head, resulting in lighter coloration than the coloring pattern commonly observed in tayras. Despite these lighter color pattern, the specimens showed parts of soft tissue, such as iris and the skin, with pigmentation very similar to that present in individuals with the typical color pattern. This set of factors indicates the specimens recorded were in fact leucistic and not albino. Among the specimens examined in the scientific collections, we found nine individuals from different localities that presented the whitish yellow color pattern. Some studies attribute the higher frequency of cases of leucism due to small populations and / or with some mechanism of reproductive isolation. Thus, analysis of the genetic variability of populations containing individuals with such characteristics should be considered. On the other hand, the occurrence of polymorphic color phenotype in tayras indicates that hypotheses related to the fixation of recessive characteristics, or on possible environmental adaptive advantages of these phenotypes can be tested.
... In contrast to all other mainland Leopard subspecies throughout the world, the Indochinese Leopard is unique in that a large percentage of remaining individuals are melanistic, including nearly 100% of the individuals in the Malay Peninsula ( Kawanishi et al. 2010). The exceptions are the populations of the Indochinese Leopard in the open dry deciduous forests of eastern and northern Cambodia, in which all individuals are spotted. ...
... The exceptions are the populations of the Indochinese Leopard in the open dry deciduous forests of eastern and northern Cambodia, in which all individuals are spotted. It has been hypothesized that melanism in the Indochinese Leopard was an adaptation to the closed canopy of tropical evergreen forests in Southeast Asia, thereby allowing better concealment of Leopard to ambush prey or avoid dominant Tiger in shadowy habitats where light seldom penetrates to the forest floor (Kawanishi et al. 2010). More recent research suggests that melanism is an adaptation in Leopard that helps with thermoregulation in hot humid forests in addition to camouflage (da Silva et al. 2017), thus there may be multiple benefits for the same gene expression. ...
... It is widely known among naturalists that " black panthers " are most frequent in southeast regions of Asia (Pocock 1929, Kingdon 1977). Kawanishi et al. (2010) confirmed general knowledge with the help of camera traps distributed in the Malay Peninsula: from 474 samples, only 29 belonged to the spotted morph and these were without any exception taken north of the Isthmus of Kra. (However, this does not mean there are no spotted individuals south of the Isthmus – Kawanishi mentions some sporadic sightings.) ...
Thesis
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The presented work investigates various theories about the significance of felid coat patterns. Most of the hypotheses are based on the theory of adaptive function of animal colouration. As regards felids, it is generally believed that the coat pattern has the function of aggressive resemblance. Early testimonies about cryptic properties of colouration of cats are summarized to provide the historical perspective of the problem. Other putative adaptive functions of patterns are investigated (intraspecific communication, thermoregulation) and close attention is paid to the option that the spotted coat of the leopard (Panthera pardus) serves as a cue for recognition from the perspective of prey. In the next stage, some of the hypotheses are tested by statistical means. The problem is simplified into search for association between a given category of a coat pattern (e.g. spots, stripes) and usual habitat of felid species (e.g. forest, grassland). Pagel’s test for correlated changes is employed to account for phylogenetic relationships. In addition to broadly acknowledged hypotheses, the validity of proposition that juvenile coat patterns have the function of protective resemblance is tested. Finally, the problem of colouration of felids is linked to broader context of theoretical biology. Apart from prevailing (neo-)Darwinian perspective to the problem of animal colouration, the theory of colour conflict by Hingston and Portmann’s aesthetical morphology are discussed at great length. It is demonstrated that Portmann’s approach yields valuable insights into the problem of the colouration of felids, mainly ontogenetic colour change. Factors accounting for marginal position of Portmann’s investigations within the framework of current biology are analysed from epistemological and philosophical perspective.
... However, it appears to occur infrequently in wild felid populations. Other somewhat familiar pelage variants in felids include both leucism (Robinson 1969, McBride & Giordano 2010) and melanism (Robinson 1970b, Wibisono & Mc-Carthy 2010, Bashir et al. 2011, Giordano et al. 2012, of which the latter variant has recently been the subject of increasing genetic and field research (Eizirik et al. 2003, Haag et al. 2010, Kawanishi et al. 2010). Although we are not aware of the existence of other fishing cat pelage variants, melanism does occur in leopard cats from the Sundarbans (Kumar Das et al. 2012). ...
... At the turn of the century, scientific interest in leopards was boosted by the introduction of long-term, well-funded, leopard research projects like The Cape Leopard Trust (www.capeleopard.org.za) and Mun-Ya- Wana Leopard Project of the Panthera Foundation (www.panthera.org). The past ten years have seen a general increase in applied ecological research along with more conservation (Martins and Martins, 2006; Aljohany, 2007; Balme et al., 2009b), behavioural (de Ruiter and Berger, 2001), evolutionary (Zuberbühler and Jenny, 2002) and genetic studies (Kawanishi et al., 2010). Concerns about human-leopard interactions and the consequences thereof have led to more publications based around carnivore conflict management and mitigation, particularly concerning non-lethal management techniques like translocation (Schiess-Meier et al., 2007; Sangay and Vernes, 2008; Weilenmann et al., 2010; Athreya et al., 2011). ...
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The African leopard Panthera pardus pardus is an apex predator, and a reliable indicator of a healthy ecosystem. Currently assessed as near threatened‘ by the IUCN Red List of Threatened Species, leopards are exposed to a range of threats that include: habitat loss and/or degradation, depletion of natural prey, conflict with people, consumptive- and non-consumptive practices, and the illegal trade of leopard products. Leopards have disappeared from 37% of their historical range, and are continuing to decline. Leopards have been extirpated across many areas of Africa, especially where people are densely populated, which has left patchily distributed leopard populations throughout the continent—to date, an accurate census has never been achieved. A variety of management techniques and conservation schemes are used in an attempt to mitigate threats directed at leopards; for example: (1) improved livestock husbandry that reduces livestock depredation and thus reduces retaliatory killing of the supposed offender—often thought to be the leopard, (2) designing community-based conservation schemes that benefit local people (i.e., food, money, jobs, ownership), (3) financial compensation for livestock losses, (4) ecotourism and regulated sport hunting, and (5) ratification of conservation-orientated government policies. Detailed, long-term leopard research began in the 1970‘s, and continues to the present day with many peer-reviewed papers focusing primarily on their ecology within savannah ecosystems. Technological advances (e.g., GPS tracking collars and remote camera traps) are helping to further our knowledge of leopard demographics, intra- and inter-specific interactions, predation, habitat use, and effective monitoring techniques. Nevertheless, more research is desperately required if leopard populations are to persist within human-dominated landscapes, like continental Africa.
... Again, southern populations are less transient, and have persisted in isolation for longer. Murie (1959) posited that the abundance of dark morphs of both Arctic and red foxes from Bering Sea and Aleutian islands are a consequence of their extended isolation; a phenomenon consistent with genetic drift within small populations, having implications for adaptive capacity under future scenarios of change (e.g., Kawanishi et al. 2010), and possibly also deterministic selection (Uy and Vargas-Castro 2015), although with unknown adaptive significance. Both Commander and Pribilof Arctic fox are considered focal populations for future conservation assessment (Geffen et al. 2007). ...
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We synthesize the historical biogeography of non-marine fauna (mammals, birds, fish, non-parasitic and parasitic invertebrates) occurring across Bering Sea islands, considering endemism, species with origins in both North America and Far East Asia, and the impacts of introduced species through the Bering region. Insular communities often constitute assemblages of unique evolutionary lineages that reflect both neutral and selective processes of diversification as a consequence of isolation and through responses to in situ environmental change. However, current knowledge of the basic distribution, ecology, and evolutionary identity of the majority of terrestrial species through the Bering Sea region is still generally lacking. A preponderance of scientific effort associated with these islands has instead focused on conservation and management of marine-associated species and economically viable biological resources. Given the critical role that terrestrial environments play in maintaining evolutionary and ecological linkages between land and sea, a greater understanding of existing biodiversity, and the biological processes that influence community integrity through this remote region is warranted. Resolving responses of resident insular species to rapidly warming Arctic climate and to modern human-associated disturbances provides valuable insight for effective management of future population trajectories and for revealing the dynamics of intra- and inter-specific connectivity across the northern hemisphere and between marine and terrestrial ecosystems. The location of this region at a high-latitude cross-roads has led to a preponderance of island taxa having trans-Beringian distributions. Species associated with the Palearctic or the Nearctic occur across a strong longitudinal gradient, reflecting the role of the Bering Sea as a dispersal filter between mainland areas. Aleutian oceanic islands reflect different biogeographic histories among taxonomic groups from land-bridge islands of the Bering Sea. We discuss evidence of recent biodiversity responses to modern environmental perturbations, including continued colonisations and novel species interactions, and call for increased scientific scrutiny of terrestrial fauna across these remote outposts.
... Several biological factors may be influenced by melanism, including thermoregulation, susceptibility or response to disease, camouflage, aposematism, sexual selection and reproductive success (Majerus 1998). Melanism is common in the Felidae, having been documented in 13 of its 38 species (Schneider et al. 2012), in some cases reaching high frequencies in natural populations (Kawanishi et al. 2010). Classical hypothesis have suggested that such coat color variants can present adaptive advantages under certain ecological conditions, but these ideas have never been rigorously tested for any wild cat. ...
... Nevertheless, poaching is presumably not as extensive as other countries in Southeast Asia, so most forested areas in the country could potentially contain leopard. Interestingly, all records from Peninsular Malaysia are of melanistic leopard (Kawanishi et al., 2010), except for a few individuals (Tan et al., 2015). ...
... This haplotype distribution indicates that after the Middle Pleistocene expansion of leopards from continental Asia to Java, a second, very recent expansion of leopards occurred from Indochina to the Malay Peninsula (Fig. 1d). Such a recent expansion and the selection of individuals with black fur may also explain the near fixation of melanism in this population (Kawanishi et al., 2010). ...
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The leopard Panthera pardus is widely distributed across Africa and Asia; however, there is a gap in its natural distribution in Southeast Asia, where it occurs on the mainland and on Java but not on the interjacent island of Sumatra. Several scenarios have been proposed to explain this distribution gap. Here, we complemented an existing dataset of 68 leopard mtDNA sequences from Africa and Asia with mtDNA sequences (NADH5 + ctrl, 724 bp) from 19 Javan leopards, and hindcasted leopard distribution to the Pleistocene to gain further insights into the evolutionary history of the Javan leopard. Our data confirmed that Javan leopards are evolutionarily distinct from other Asian leopards, and that they have been present on Java since the Middle Pleistocene. Species distribution projections suggest that Java was likely colonized via a Malaya-Java land bridge that by-passed Sumatra, as suitable conditions for leopards during Pleistocene glacial periods were restricted to northern and western Sumatra. As fossil evidence supports the presence of leopards on Sumatra at the beginning of the Late Pleistocene, our projections are consistent with a scenario involving the extinction of leopards on Sumatra as a consequence of the Toba super volcanic eruption (~74 kya). The impact of this eruption was minor on Java, suggesting that leopards managed to survive here. Currently, only a few hundred leopards still live in the wild and only about 50 are managed in captivity. Therefore, this unique and distinctive subspecies requires urgent, concerted conservation efforts, integrating in situ and ex situ conservation management activities in a One Plan Approach to species conservation management.
... A variação da tonalidade também pode estar associada à distribuição geográfica, variação sazonal, idade, sexo e peso (Ortolani, 1999;Stoner et al., 2003;Kawanishi et al., 2010;Ancilloto e Mori, 2017). No entanto, anomalias de cores são observadas quando pigmentos tegumentares estão presentes em quantidades excessivas ou deficientes em partes ou na totalidade do corpo (Prota, 1980;Ortolani, 1999;Fertl e Rosel, 2002;Caro, 2005), sendo estas descritas como leucismo, albinismo e melanismo (Fertl e Rosel, 2002). ...
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RESUMO-Mamíferos de coloração anômala ocorrem por distúrbios genéticos, sendo um evento não habitual na natureza, porém relatado em várias espécies. Este trabalho apresenta a ocorrência de Eira barbara com coloração anômala, consideradas leucísticas. O registro foi realizado por captura de imagem em armadilha fotográfica na região central do estado de São Paulo, no município de Jaú, sendo o primeiro relato em remanescente de Floresta Estacional Semidecidual no Brasil. Palavras-chave: Mustelidae; Distúrbio genético; Pelagem anômala; Leucismo. ABSTRACT-Anomalous colored mammals occur due to genetic disorders, being an unusual event in nature. This work present the occurrence of Eira barbara with anomalous coloring, considered leucistic. The recording was performed by image capture in a camera trap in the central region of the state of São Paulo, in the municipality of Jau, being the first report in remnants of Seasonal Semideciduos Forest in Brazil.
... The biological significance and even the geographical distribution of felid melanistic variants are still poorly understood, but recent analyses indicate that melanism in some cases can reach very high frequencies in regional populations (e.g. Kawanishi et al., 2010;Hedges et al., 2015). Such observations support the hypothesis that melanism can provide an adaptive advantage in certain ecological conditions Caro, 2005;Allen et al., 2011). ...
... Nevertheless, we considered most forested areas outside of PAs as potential, unless leopard was not detected in an area, given that poaching is presumably not as extensive as in other countries of Southeast Asia. All records from Peninsular Malaysia are of melanistic leopard (Kawanishi et al., 2010), except for a few individuals (Tan et al., 2015). The total area of confirmed and potential distribution is 47,051 km 2 , and the population estimate is 282-847 total individuals, with 119-356 breeding adults. ...
Article
The Indochinese leopard (Panthera pardus delacouri) is a genetically distinct subspecies that historically occurred throughout mainland Southeast Asia, but might have experienced recent declines in numbers and distribution. This study aimed to determine the current distribution of the Indochinese leopard, and estimate its population size, by reviewing data from camera trap and other wildlife surveys conducted during the past 20 years. Our results showed the Indochinese leopard likely now occurs only in 6.2% of its historical range, with only 2.4% of its distribution in areas of confirmed leopard presence. The leopard is extirpated in Singapore, likely extirpated in Laos and Vietnam, nearly extirpated in Cambodia and China, and has greatly reduced distributions in Malaysia, Myanmar, and Thailand. There are plausibly only two major strongholds remaining, which we consider priority sites: Peninsular Malaysia, and the Northern Tenasserim Forest Complex. We also identified a small isolated population in eastern Cambodia as a third priority site, because of its uniqueness and high conservation value. We estimate a total remaining population of 973–2503 individuals, with only 409–1051 breeding adults. Increased poaching for the illegal wildlife trade likely is the main factor causing the decline of the Indochinese leopard. Other potential contributing factors include prey declines, habitat destruction, and possibly disease. We recommend a separate IUCN assessment for the Indochinese leopard, and that this subspecies be classified as Endangered. Our findings provide important information that can help guide where conservation actions would be most effective in preventing the extinction of this subspecies.
... The biological significance and even the geographical distribution of felid melanistic variants are still poorly understood, but recent analyses indicate that melanism in some cases can reach very high frequencies in regional populations (e.g. Kawanishi et al., 2010;Hedges et al., 2015). Such observations support the hypothesis that melanism can provide an adaptive advantage in certain ecological conditions Caro, 2005;Allen et al., 2011). ...
... Felid melanism is especially interesting because it is polymorphic within each species, but its possible adaptive significance has received little attention to date [9]. In rodents and in birds, an adaptive role of melanism is well-established from ecological and field studies [1,2,10,11], but the evolutionary forces that underlie melanism in larger mammals are less clear, with examples of introgression in the case of North American black wolves [12], and a founder effect in the case of Malaysian black leopards [13]. A recent study by Allen et al. [9] found that the presence of melanism across felid species is correlated with habitat and/or behavioral diversity, and suggested a common underlying mechanism of disruptive selection. ...
... These findings further supported the inference that melanism arose multiple times in the Felidae and reached high population-level frequencies in some areas, albeit never achieving complete fixation species wide. A striking case were the leopards from the Malay Peninsula in Southeast Asia, which exhibit almost complete regional fixation for melanism (76). This intriguing observation was explored in more detail by systematically surveying and mapping the geographic distribution of leopard melanism across the species' range, which demonstrated that this phenotype is significantly associated with tropical moist forests and significantly absent from drier, open habitats, such as xeric scrublands (77). ...
Article
The diversity of mammalian coat colors, and their potential adaptive significance, have long fascinated scientists as well as the general public. The recent decades have seen substantial improvement in our understanding of their genetic bases and evolutionary relevance, revealing novel insights into the complex interplay of forces that influence these phenotypes. At the same time, many aspects remain poorly known, hampering a comprehensive understanding of these phenomena. Here we review the current state of this field and indicate topics that should be the focus of additional research. We devote particular attention to two aspects of mammalian pigmentation, melanism and pattern formation, highlighting recent advances and outstanding challenges, and proposing novel syntheses of available information. For both specific areas, and for pigmentation in general, we attempt to lay out recommendations for establishing novel model systems and integrated research programs that target the genetics and evolution of these phenotypes throughout the Mammalia. Expected final online publication date for the Annual Review of Animal Biosciences, Volume 9 is February 16, 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
... Despite the vital role of colouration for species, individuals of unusual colours can arise in a population. If these atypical coloured specimens perform in the same way or better than the regular coloured individuals, the new colouration may be retained in the population for several generations and become relatively common, such as in the melanistic wolves Canis lupus of western North America (Musiani et al. 2007) and leopards Panthera pardus in south-east Asia (Kawanishi et al. 2010). ...
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Records of atypical black individuals of Neotropical canids are extremely rare. Here, we report the first ever record of a black-coloured maned wolf Chrysocyon brachyurus, either wild or in captivity. Using camera traps to survey a protected area in the Brazilian Cerrado we obtained 30 maned wolf photographic records, 25 in its common reddish colour and five (16.6% of the records) in the previously unknown black phenotype. We suggest this is possibly an independent event of melanism in canids, discuss its potential evolutionary benefits and give recommendations for further research.
... Melanism in golden cats has �een traced to a mutation in the coding region of the Agouti Signaling Protein ASIP, leading to a failure in producing light coat pigmentation and a pro� duction in eumelanin (Schneider et al. 2012). This mutation is thought to follow recessive inheritance patterns and, in other felid spe� cies, has nearly reached fixation (Kawanishi et al. 2010), suggesting an adaptive �enefit gained from melanism. There is evidence that in some areas of the golden cat range, such as Sikkim, India, where all recorded individu� als have �een melanistic (Bashier et al. 2011) and at higher elevations in Bhutan (Jigme 2011) where multiple melanistic individuals have �een recorded, melanism may also con� fer an adaptive �enefit. ...
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Asiatic golden cats Catopuma temminckii are currently listed as Near Threatened in the IUCN Red List with a declining population trend, largely due to habitat loss and poaching. As one of the most polymorphic felids, multiple pelage morphs have been recorded in the northern parts of its range, while in the southern parts, the reddish-brown coat is dominant. In central Sumatra, field teams conducted an ad-hoc camera trap study in Rimbang Baling Wildlife Reserve from March - June 2016. Seven videos of golden cats were recorded, including one melanistic individual. Although camera trapping efforts have taken place throughout Sumatra for years, this is only the second recorded melanistic individual, suggesting rare occurrence of the recessive mutation leading to melanism in Sumatra, despite more common occurrence elsewhere.
... In general, among large cats, melanism is usually more common in those populations that live in dense forests. With the reduced light, dark animals are less noticeable in such environments compared to open areas, giving them a selective advantage in hunting and survival (Kawanishi et al., 2010). A high frequency of occurrence of melanists is also noted among some species of pikas and hares. ...
Article
Cases of melanism in the mountain hare Lepus timidus in the territory of Yakutia were analyzed. The highest frequency of this rare phenomenon was observed in the basin of the Vilyuy River, with at least nine cases over the last 50 years. It was proposed that this phenomenon was the result of increased mutagenesis in this territory, due to consequence of the features of the natural geochemical background and anthropogenic pollution.
... We acknowledge several caveats in our study. Despite our attempt to obtain as complete a set of published studies for our analysis as possible, at least three papers with nine additional records of leopard [81,82] and one record of a flat-headed cat [83] escaped our attention. Of these 10 records, including 29 recently released records of threatened and near-threatened carnivores [84][85][86][87], 82% occurred within the hotspots identified in our analysis, confirming the importance of these regions for carnivore conservation. ...
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Mammalian carnivores play a vital role in ecosystem functioning. However, they are prone to extinction because of low population densities and growth rates, and high levels of persecution or exploitation. In tropical biodiversity hotspots such as Peninsular Malaysia, rapid conversion of natural habitats threatens the persistence of this vulnerable group of animals. Here, we carried out the first comprehensive literature review on 31 carnivore species reported to occur in Peninsular Malaysia and updated their probable distribution. We georeferenced 375 observations of 28 species of carnivore from 89 unique geographic locations using records spanning 1948 to 2014. Using the Getis-Ord Gi*statistic and weighted survey records by IUCN Red List status, we identified hotspots of species that were of conservation concern and built regression models to identify environmental and anthropogenic landscape factors associated with Getis-Ord Gi* z scores. Our analyses identified two carnivore hotspots that were spatially concordant with two of the peninsula's largest and most contiguous forest complexes, associated with Taman Negara National Park and Royal Belum State Park. A cold spot overlapped with the southwestern region of the Peninsula, reflecting the disappearance of carnivores with higher conservation rankings from increasingly fragmented natural habitats. Getis-Ord Gi* z scores were negatively associated with elevation, and positively associated with the proportion of natural land cover and distance from the capital city. Malaysia contains some of the world's most diverse carnivore assemblages, but recent rates of forest loss are some of the highest in the world. Reducing poaching and maintaining large, contiguous tracts of lowland forests will be crucial, not only for the persistence of threatened carnivores, but for many mammalian species in general.
... Observations of black leopards (Panthera pardus) primarily come from Southeast Asia, with few observations in Africa (da Silva et al., 2017;Sunquist & Sunquist, 2002). In the most extreme case, leopards in Peninsular Malaysia show melanism nearly to fixation in the population (Kawanishi et al., 2010). Melanism in leopards is associated with a mutation to the Agouti Signaling Protein gene that causes a loss of the normal function (Eizirik et al., 2003;Schneider et al., 2012Schneider et al., , 2015 and is recessive in its inheritance (Robinson, 1969(Robinson, , 1970. ...
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Observations of black leopards (Panthera pardus) primarily come from Southeast Asia, with few observations in Africa (da Silva et al., 2017; Sunquist & Sunquist, 2002). In the most extreme case, leopards in Peninsular Malaysia show melanism nearly to fixation in the population (Kawanishi et al., 2010). Melanism in leopards is associated with a mutation to the Agouti Signaling Protein gene that causes a loss of the normal function (Eizirik et al., 2003; Schneider et al., 2012, 2015) and is recessive in its inheritance (Robinson, 1969, 1970). The adaptive significance of melanism is not clearly understood, but most hypotheses suggest environment as a potential driver (Eizirik et al., 2003; Ortolani & Caro, 1996), with higher frequencies of black leopards found in tropical and humid environments (da Silva et al., 2017). Gathering a broad array of melanistic leopard observations, da Silva et al. (2017) reported five sightings of black leopards in Africa (P. p. pardus), but could confirm only one. The confirmed report was from Addis Ababa, Abissynya, Ethiopia, in 1909, in the form of a photograph stored at the National Museum of Natural History in the United States (da Silva et al., 2017). In Kenya, reports of black leopard are known (da Silva et al., 2017; Sunquist & Sunquist, 2002), but none have been accompanied with photographic evidence. Here, we provide photographic evidence for the existence of black leopard in Laikipia County, Kenya. We also compare the habitat of these sightings to the expectations of melanism as an adaptive trait driven by environment.
... Leopard population in Southeast Asia is found in Myanmar, Thailand, Vietnam, island of Java and Peninsular Malaysia. Interestingly, all leopard individuals recorded in Peninsular Malaysia have been of melanistic type, except for few individuals (Kawanishi et al., 2010;Tan et al., 2015). Leopard population in Peninsular Malaysia was assessed as endangered (DWNP, 2017). ...
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Aim: This research assessed the distribution of leopard to predict the habitat suitability in Taman Negara National Park and adjacent forest area. Methodology: Environmental factors for habitat suitability were derived from geographical information system (GIS) data such as elevation, slope, land-use, distance from urban and distance from river. Leopard presence data from 1993 to 2008 were integrated with the environmental parameters using maximum entropy (MaxEnt) modeling to assess habitat suitability across the study area. Results: The results showed that distance from river contributed the most (39.3%) in the habitat suitability modeling followed by distance from urban (31.4%), elevation (12.3%), land use types (10.1%), and slope (6.9%). Distance from river and urban showed highest contribution that influenced leopard distribution in which most suitable habitat occurred in proximity with river and further from urban. Habitat suitability of leopard were distributed among 48% over 2,218,389 ha of the study area. Interpretation: The findings of this study provides knowledge on how the species move and exploit different habitat niches for more effective conservation management. It provide models for future wildlife conservation and urban planning.
... While these subspecific designations likely overestimated the actual number of genetically distinct leopard lineages, it is worth noting that descriptive characteristics (e.g. pelage morphology) are often associated with unique geographic features including the near fixation of melanism in Asiatic leopards south of the Isthmus of Kra (Kawanishi et al. 2010). With the advent and application of genetic analyses, we are better able to evaluate the taxonomic hypotheses proposed by previous researchers within the context of phylogeography (Avise 2000). ...
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Once found throughout Africa and Eurasia, the leopard (Panthera pardus) was recently uplisted from Near Threatened to Vulnerable by the International Union for the Conservation of Nature (IUCN). Historically, more than 50% of the leopard's global range occurred in continental Africa, yet sampling from this part of the species' distribution is only sparsely represented in prior studies examining patterns of genetic variation at the continental or global level. Broad sampling to determine baseline patterns of genetic variation throughout the leopard's historical distribution is important, as these measures are currently used by the IUCN to direct conservation priorities and management plans. By including data from 182 historical museum specimens, faecal samples from ongoing field surveys, and published sequences representing sub-Saharan Africa, we identify previously unrecognized genetic diversity in African leopards. Our mtDNA data indicates high levels of divergence among regional populations and strongly differentiated lineages in West Africa on par with recent studies of other large vertebrates. We provide a reference benchmark of genetic diversity in African leopards against which future monitoring can be compared. These findings emphasize the utility of historical museum collections in understanding the processes that shape present biodiversity. Additionally, we suggest future research to clarify African leopard taxonomy and to differentiate between delineated units requiring monitoring or conservation action.
... It is the most widespread felid, extend� ing across much of Africa, and Asia from the Middle East to the Pacific Ocean, encompass� ing a diverse array of ha�itats, from deserts to rainforests, and from the humid tropics to temperate zones (Jaco�son et al. 2016). Me� lanism has �een long known to occur in this species across its range and is reported at very high frequencies in some areas of South� east Asia (Kawanishi et al. 2010, Clements 2015, Hedges et al. 2015 Sterndale 1984, Daniel 1996, da Silva et al. 2017. However, the only reports of melanis� tic leopards from Odisha were unconfirmed (Anonymous 1999). ...
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Although melanistic leopards Panthera pardus have been frequently observed in parts of South Asia and occasionally in India, no such records exist for Odisha. During camera trap surveys of large carnivores in Sundargarh Forest Division, Odisha, India, we obtained photographs of melanistic leopards between January 2018 and April 2018. These are the first confirmed records of melanistic leopard in Odisha, eastern India.
... Capture-recapture methods have been widely used to estimate abundance and density from camera photos in many carnivores, such as in the snow leopard (P. uncia), tiger [13][14][15], bobcat (Lynx rufus), (Alonso et al. 2015), black bear (Ursus americanus) (Fusaro et al. 2017), jaguar (P. onca) and common leopard (P. ...
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Visual leopard identifications performed with camera traps using the capture–recapture method only consider areas of the skin that are visible to the equipment. The method presented here considered the spot or rosette formations of either the two flanks or the face, and the captured images were then compared and matched with available photographs. Leopards were classified as new individuals if no matches were found in the existing set of photos. It was previously assumed that an individual leopard’s spot or rosette pattern would not change. We established that the spot and rosette patterns change over time and that these changes are the result of injuries in certain cases. When compared to the original patterns, the number of spots may be lost or reduced, and some spots or patterns may change in terms of their prominence, shape, and size. We called these changes “obliterate changes” and “rejig changes”, respectively. The implementation of an earlier method resulted in a duplication of leopard counts, achieving an error rate of more than 15% in the population at Yala National Park. The same leopard could be misidentified and counted multiple times, causing overestimated populations. To address this issue, we created a new two-step methodology for identifying Sri Lankan leopards. The multi-point identification method requires the evaluation of at least 9–10 spot areas before a leopard can be identified. Moreover, the minimum leopard population at the YNP 1 comprises at least 77 leopards and has a density of 0.5461 leopards per km2.
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Most species of wild felids are threatened, but for many little is known about their status in the wild. For the cryptic and elusive Vulnerable Sunda clouded leopard Neofelis diardi , key metrics such as abundance and occupancy have been challenging to obtain. We conducted an intensive survey for this species on the Indonesian island of Sumatra. We deployed camera traps across four study areas that varied in elevation and threats, for a total of 28,404 trap nights, resulting in 114 independent clouded leopard photographs, in which we identified 18 individuals. Using a Bayesian spatially explicit capture–recapture analysis, we estimated clouded leopard density to be 0.8–2.4 individuals/100 km ² . The highest predicted occurrence of people was at lower altitudes and closer to the forest edge, where we categorized more than two-thirds of people recorded by camera traps as bird poachers, 12.5% each as ungulate/tiger poachers and non-timber collectors, and < 2% as fishers. Our findings provide important insights into the status of this little known species in Sumatra. We recommend that the large volume of camera-trap data from other Sumatran landscapes be used for an island-wide assessment of the clouded leopard population, to provide up-to-date and reliable information for guiding future conservation planning.
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Managing populations of large carnivore species that are threatened yet in competition with each other presents a conservation challenge over which species to prioritise. We investigated interspecific competition between the Endangered tiger and two sympatric carnivore species (Near-Threatened leopard and Endangered dhole) in Malaysia. We used a spatially-explicit camera trapping method to sample the three carnivore species in order to assess their: relative abundance; spatial co-occurrence; and, temporal overlap. Our focal landscape consists of two adjacent study areas that have different forest management regimes: 1) Royal Belum State Park (RBSP) which has relatively higher levels of enforcement resulting in high tiger and prey densities, and 2) Temengor Forest Reserve (TFR) which has no enforcement resulting in low tiger and prey densities. Our analyses found interspecific competition in the form of fine-scale spatial avoidance of tiger by leopard in RBSP, but not in TFR. Thus, leopard was less likely to occur in forest patches with greater tiger presence in RBSP. Dhole was not detected in RBSP, and showed no fine-scale spatial avoidance of tiger on TRF. It was unclear if absence of dhole in RBSP was due to the high tiger density, or another factor such as disease. Regardless, our results indicate dhole can persist in human-degraded forest landscapes. Neither leopards nor dholes showed any major temporal avoidance of tigers on either site. Our findings of fine-scale spatial avoidance by leopard towards tiger and possible exclusion of dhole by tiger in areas of high tiger density were similar to that found in other areas of Asia. Our study demonstrates that degraded forest can be important for threatened carnivores. It also provides a reminder that efforts to recover tiger populations, a conservation priority, may require flanking measures to secure meso-predator populations, especially those forced into edge environments where they come into greater contact with people.
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Melanism is a form of pigmentation polymorphism where individuals have darker colouration than what is considered the 'wild' phenotype. In the case of leopards, Panthera pardus, melanism occurs at higher frequencies amongst populations in tropical and subtropical moist forests of south and southeast Asia, presenting a unique challenge in estimating and monitoring these populations. Unlike the wild phenotype that is readily recognizable by its rosette patterns, melanism results in individuals being unidentifiable or 'unmarked' through photographic captures obtained using white flash cameras. Spatial mark-resight (SMR) models that require only a subset of the population to be 'marked' offer the opportunity to estimate population density. In this study, we present an application of SMR models to estimate leopard densities using camera trap survey data from three sampling years at Manas National Park (MNP), India. By using an SMR model that allowed us to include captures of unidentified sightings of marked individuals, we were also able to incorporate captures where identity was either not confirmed or only known from a single flank. Following 18 674 trap-days of sampling across three years, we obtained 728 leopard photo-captures, of which 22.6% (165) were melanistic. We estimated leopard densities of 4.33, 2.61 and 3.37 individuals/100 km 2 across the 3 years. To our best knowledge, these represent the first known estimates of leopard densities from populations comprising both melanistic and wild phenotypes. Finally, we highlight that SMR models present an opportunity to revisit past camera trap survey data for leopards and other species such as Jaguars, P. onca, that exhibit phenotypic polymorphism towards generating valuable information on populations.
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Some wild felines have a diverse range of coat colors while others do not. Jaguars and leopards, for instance, come in spotted and melanistic forms but tigers are always striped and lions always beige. Smaller cats like clouded leopards, marbled cats, and ocelots are almost always patterned in the same way while jaguarundis, oncillas, and golden cats occur in several different colors and patterns. This article is protected by copyright. All rights reserved.
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Mammalian colors and color patterns are some of the most diverse and conspicuous traits found in nature and have been widely studied from genetic/developmental and evolutionary perspectives. In this review we first discuss the proximate causes underlying variation in pigment type (i.e., color) and pigment distribution (i.e., color pattern) and highlight both processes as having a distinct developmental basis. Then, using multiple examples, we discuss ultimate factors that have driven the evolution of coloration differences in mammals, which include background matching, intra- and interspecific signaling, and physiological influences. Throughout, we outline bridges between developmental and functional investigatory approaches that help broaden knowledge of mammals' memorable external appearances, and we point out areas for future interdisciplinary research.
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Morphological variation in natural populations is a genomic test bed for studying the interface between molecular evolution and population genetics, but some of the most interesting questions involve non-model organisms that lack well annotated reference genomes. Many felid species exhibit polymorphism for melanism but the relative roles played by genetic drift, natural selection, and interspecies hybridization remain uncertain. We identify mutations of Agouti signaling protein (ASIP) or the Melanocortin 1 receptor (MC1R) as independent causes of melanism in three closely related South American species: the pampas cat (Leopardus colocolo), the kodkod (Leopardus guigna), and Geoffroy's cat (Leopardus geoffroyi). To assess population level variation in the regions surrounding the causative mutations we apply genomic resources from the domestic cat to carry out clone-based capture and targeted resequencing of 299 kb and 251 kb segments that contain ASIP and MC1R, respectively, from 54 individuals (13-21 per species), achieving enrichment of ~500-2500-fold and ~150x coverage. Our analysis points to unique evolutionary histories for each of the three species, with a strong selective sweep in the pampas cat, a distinctive but short melanism-specific haplotype in the Geoffroy's cat, and reduced nucleotide diversity for both ancestral and melanism-bearing chromosomes in the kodkod. These results reveal an important role for natural selection in a trait of longstanding interest to ecologists, geneticists, and the lay community, and provide a platform for comparative studies of morphological variation in other natural populations.
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Historically, melanistic fox squirrels have been found in Council Bluffs, Iowa, and along the Missouri River in Eastern Omaha, Nebraska. However, recent anecdotal observations suggest that the melanistic trait in fox squirrels is expanding westward into Omaha. Squirrels were surveyed in the autumn of 2010 and 2011 along transect lines within five major city sections and different habitat types; comparisons were made to a survey performed in 1973. Proportion of melanistic squirrels in Council Bluffs remained at levels similar to 1973 surveys (∼50% melanistic). Since 1973 melanistic fox squirrels have increased in Omaha, with a higher proportion of melanistic squirrels in northwest (7.4%) and northeast (7.6%) compared to southwest or southeast Omaha (4.6%). Melanistic squirrels were found in higher proportions in parks (12.1%) and residential (10.9%) habitats compared to business (6.1%), industrial (7.4%), or golf course (8.0%) habitats. Melanistic squirrels were also observed more frequently at colder temperatures than rufus squirrels. The results yielded significant variation in the percent of melanistic individuals in each section and suggest the proportion of melanistic individuals is increasing westward.
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Melanistic fox squirrels (Sciurus niger) have expanded westward and increased in frequency in the Omaha, Nebraska, and Council Bluffs, Iowa, metropolitan areas. The selective advantage of melanism is currently unknown, but thermal advantages have been hypothesized, especially in winter. No difference in metabolic response curves were measured between melanistic (black) and rufus (orange) fox squirrels. When exposed to sunny skies, both melanistic and rufus squirrels had higher surface (skin and fur) temperature as ambient temperatures increased. Melanistic squirrel surface temperatures did not differ when squirrels were exposed to sunny or cloudy skies. However, rufus individuals showed significantly lower increases in surface temperatures when under cloudy skies. During fall months, rufus individuals were about 1.5 times more active throughout the day than melanistic individuals. However, in winter, melanistic fox squirrels were approximately 30% more active in the mornings (before 13:00) compared to rufus squirrels. Pre-winter body condition was higher in melanistic (25.5 ± 1.8 g/cm) compared to rufus (20.30 ± 3.6 g/cm) fox squirrels; however, there were no significant differences between melanistic (22.8 ± 1.4 g/cm) and rufus (23.9 ± 0.8 g/cm) fox squirrel post-winter body condition. The results of this study indicate that melanistic fox squirrels may have a slight winter thermal advantage over rufus fox squirrels by maintaining higher skin temperatures.
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ACKNOWLEDGMENTS The success and completion of this 6-year University of Florida-Malaysia Tiger Project are due to support from many individuals and organizations. Foremost, I would like to thank my major professor, Dr. Mel Sunquist, and his partner, Fiona Sunquist, for their tireless guidance and encouragement. His generosity and courage to take on foreign students who always require more attentions and logistics deserve honor. Had it not been for the serendipity to have become acquainted with Fiona, none of this would have been realized. Fiona’s trademark feel-good-talks often helped me lift sunken spirits. I would also like to thank my committee members, Dr. John Eisenberg, Dr. Mike Moulton, Dr. Jim Nichols, and Dr. Scot Smith, for critiques and comments on the dissertation. Dr Nichols trained me on the science of population estimation. His patience with me was much appreciated. Although not as an official committee member, advice and support provided by Dr. Ullas Karanth of the Wildlife Conservation Society-India were vital to the project. My intellectual phenology,has been shaped by the works of Dr. Eisenberg and Dr. Larry Harris, under whom I was fortunate to learn during their last years at the University of Florida. My spirit for the carnivore conservation was fueled by the works and characters of Drs. Mel Sunquist, Ullas Karanth, Howard Quigley, Maurice Hornocker, Alan Rabinowitz, and Dale Miquelle, who have provided me with training and opportunities at some point in my life. Lastly in the Department, the additional work by Cynthia Sain, Caprice McRay, and Monica Lindberg was important for me to maintain UF student status during my absence. Financial support for the project was provided by the Save the Tiger Fund, a special project of the National Fish and Wildlife Foundation created in partnership with ExxonMobil Corporation, University of Florida, Disney Wildlife Conservation Fund, World Wide Fund for iii Nature (WWF)-Japan, WWF-UK, WWF-Netherlands,and 21,Century Tiger- a partnership between Global Tiger Patrol and London Zoo. Other support from WWF-Malaysia, Wildlife
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We sampled riparian frogs along 18 streams at eight localities in Borneo. At four of these sites we sampled during more than one year. Altogether 49 species were included in our study and total sample size was 13,249. We measured overlap in species occurrences and arrays of abundances within and among localities. Variation over the time span of our study was minor within communities. Overlaps between streams at a locality were generally higher than overlaps of pairs of streams from different localities. Environmental variation, particularly in stream width and gradient, had a clear effect on both intra-and inter-locality overlaps. Although rainfall varied between localities and within localities over time, that variation did not seem to affect overlaps among or within communities. Environmental factors did not account for all differences in overlaps between communities. Instead, regional processes, perhaps the timing of barriers or speciation events, appear to have been responsible for geographic restrictions of several species, leading to variation in overlap values.
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The monitoring and management of species depends on reliable population estimates, and this can be both difficult and very costly for cryptic large vertebrates that live in forested habitats. Recently developed camera trapping techniques have already been shown to be an effective means of making mark-recapture estimates of individually identifiable animals (e.g. tigers). Camera traps also provide a new method for surveying animal abundance. Through computer simulations, and an analysis of the rates of camera trap capture from 19 studies of tigers across the species' range, we show that the number of camera days/tiger photograph correlates with independent estimates of tiger density. This statistic does not rely on individual identity and is particularly useful for estimating the population density of species that are not individually identifiable. Finally, we used the comparison between observed trapping rates and the computer simulations to estimate the minimum effort required to determine that tigers, or other species, do not exist in an area, a measure that is critical for conservation planning.
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Color maps are presented that depict Holocene sea level transgressions and probable submerged lakes on the Sunda Shelf in Southeast Asia. The present-day topography of the Sunda Shelf and the existence of present-day submerged depressions on the Sunda Shelf have been detected through spatial analysis of the Sunda Shelf using the Digital Elevation Model (DEM) developed from the ETOPO2 Global 2' Elevation data. These depressions could be paleo-lakes that existed when the Sunda Shelf was exposed during the Last Glacial Maximum (LGM). These depressions were gradually submerged when the sea level began to rise from –116 m below present-day levels (BPL) during the terminal phase of the LGM, 21 thousand years before present (ka BP) to its maximum, +5m above present-day mean sea level (MSL), during the mid-Holocene (4.2 ka BP). The topography of these depressions is presented on maps and discussed. The results of this study identify several large submerged depressions that may contain sediment dated from the LGM to the mid-Holocene and we recommend that sediment layers be sampled to confirm or disprove the presence of these proposed fresh water paleo-lakes on the Sunda Shelf.
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Aim To describe current geographical patterns of genetic diversity and infer the his-torical population dynamics of the stone oaks (Lithocarpus) in Southeast Asia. Location We sampled three populations in Indochina: (1) Yunnan province, China; (2) Pyin Oo Lwin area, Myanmar and (3) north-western Vietnam; two in western Borneo: (1) South-western Sarawak and (2) West Kalimantan, Indonesia; two in central Borneo: (1) north coastal Sarawak and (2) north-eastern Sarawak, Malaysia; and two in northern Borneo: (1) Central Sabah and (2) Northern Sabah, Malaysia. Methods A phylogenetic reconstruction of chloroplast DNA sequence variation from numerous individuals of multiple species was used to determine geographical distribu-tion of genetic diversity. A resampling scheme was used to determine the significance of these patterns at different hierarchical levels of the phylogeny. Results were compared with a previously published set of nuclear DNA sequence data.
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Aim We describe the distributions of mammal species between the Indochinese and Sundaic subregions and examine the traditional view that the two faunas show a transition near the Isthmus of Kra on the Thai–Malay peninsula. Location Species distributions are described along a 2000-km transect from 20° N (northernmost Thailand) to 1° N (Singapore). Methods For the 325 species of native non-marine mammals occurring along the transect we used published records to provide a database of their distributional records by degree of latitude. Results Along the transect we found 128 Indochinese species with southern range limits, 121 Sundaic species with northern range limits, four un-assignable endemics and 72 widespread species. In total, 152 southern and 147 northern range limits were identified, and their distribution provides no evidence for a narrow faunal transition near the Isthmus of Kra (10°30′ N) or elsewhere. Range limits of both bats and non-volant mammals cluster in northernmost peninsular Malaysia (5° N) and 800 km further north, where the peninsula joins the continent proper (14° N). The clusters of northern and southern range limits are not concordant but overlap by 100–200 km. Similarly, the range limits of bats and non-volant mammals cluster at slightly different latitudes. There are 30% fewer species and range limits in the central and northern peninsula (between 6 and 13° N), and 35 more widely distributed species have range gaps in this region. In addition, we found 70 fewer species at the southern tip of the peninsula (1° N) than at 3–4° N. Main conclusions The deficiencies of both species and species range limits in the central and northern peninsula are attributed to an area effect caused by repeated sea-level changes. Using a new global glacioeustatic curve developed by Miller and associates we show that there were > 58 rapid sea-level rises of > 40 m in the last 5 Myr that would have resulted in significant faunal compression and local population extirpation in the narrow central and northern parts of the peninsula. This new global sea-level curve appears to account for the observed patterns of the latitudinal diversity of mammal species, the concentration of species range limits north and south of this area, the nature and position of the transition between biogeographical subregions, and possibly the divergence of the faunas themselves during the Neogene. The decline of species diversity at the southern end of the transect is attributed to a peninsula effect similar to that described elsewhere.
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A comprehensive survey on large mammal diversity from a disturbed forest in Peninsular Malaysia has been carried out for over a period of 21months. A total of 24 camera traps which accumulated to 5972trap days. A total of 33 species 27 genera and 15 families of mammals were recorded via camera trapping and observations. The use of camera traps provides detailed information on diversity of some cryptic and secretive mammals. Secondary forest may support a wide diversity of mammals at a stable condition where intrusion, excision and fragmentation are reduced or avoided. The threats to mammals in the study are also discussed.
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This study provided the first reliable density estimate of tigers based on photographic capture data in Taman Negara National Park, Peninsular Malaysia's most important conservation area. Estimated densities () of adult tigers ranged from 1.10 ± 0.52 to 1.98 ± 0.54 tigers/100 km2 (X2=1.56, df=2, P=0.46) with the overall mean of 1.66 ± 0.21 tigers/100 km2. The tiger population in the 4343-km2 park was estimated to be 68 (95% CI: 52–84) adult tigers. Prey biomass estimates ranged from 266 to 426 kg/km2, and wild boar were the most important potential prey species in terms of abundance, biomass, and occupancy, followed by muntjac. Both tigers and leopards were more diurnal than nocturnal, which corresponded with the activity patterns of wild boar and muntjac. No evidence of poaching of large mammals was found in the 600-km2 study sites and overall human impacts on the tiger–prey community appear to be minimal, but in the long run its viability needs to be evaluated in a greater landscape context.
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Eight traditional subspecies of tiger (Panthera tigris),of which three recently became extinct, are commonly recognized on the basis of geographic isolation and morphological characteristics. To investigate the species' evolutionary history and to establish objective methods for subspecies recognition, voucher specimens of blood, skin, hair, and/or skin biopsies from 134 tigers with verified geographic origins or heritage across the whole distribution range were examined for three molecular markers: (1) 4.0 kb of mitochondrial DNA (mtDNA) sequence; (2) allele variation in the nuclear major histocompatibility complex class II DRB gene; and (3) composite nuclear microsatellite genotypes based on 30 loci. Relatively low genetic variation with mtDNA,DRB,and microsatellite loci was found, but significant population subdivision was nonetheless apparent among five living subspecies. In addition, a distinct partition of the Indochinese subspecies P. t. corbetti in to northern Indochinese and Malayan Peninsula populations was discovered. Population genetic structure would suggest recognition of six taxonomic units or subspecies: (1) Amur tiger P. t. altaica; (2) northern Indochinese tiger P. t. corbetti; (3) South China tiger P. t. amoyensis; (4) Malayan tiger P. t. jacksoni, named for the tiger conservationist Peter Jackson; (5) Sumatran tiger P. t. sumatrae; and (6) Bengal tiger P. t. tigris. The proposed South China tiger lineage is tentative due to limited sampling. The age of the most recent common ancestor for tiger mtDNA was estimated to be 72,000-108,000 y, relatively younger than some other Panthera species. A combination of population expansions, reduced gene flow, and genetic drift following the last genetic diminution, and the recent anthropogenic range contraction, have led to the distinct genetic partitions. These results provide an explicit basis for subspecies recognition and will lead to the improved management and conservation of these recently isolated but distinct geographic populations of tigers.
Article
With the increasing popularity of remote photography in wildlife research, a large variety of equipment and methods is available to researchers. To evaluate advantages and disadvantages of using various types of equipment for different study objectives, we reviewed 107 papers that used either time-lapse or animal-triggered photography to study vertebrates in the field. Remote photography was used primarily to study avian nest predation, feeding ecology, and nesting behavior; additional applications included determining activity patterns, presence-absence monitoring, and estimating population parameters. Using time-lapse equipment is most appropriate when animals occur frequently in the photographic frame, the activity of interest occurs repeatedly, or no distinct event occurs to trigger a camera. In contrast, animal-triggered (light or mechanically triggered) systems are appropriate when events occur infrequently or unpredictably and there is a great likelihood that a trigger will be activated. Remote photography can be less time consuming, costly, and invasive than traditional research methods for many applications. However, researchers should be prepared to invest time and money troubleshooting problems with remote camera equipment, be aware of potential effects of equipment on animal behavior, and recognize the limitations of data collected with remote photography equipment.
Article
This study provided the first reliable density estimate of tigers based on photographic capture data in Taman Negara National Park, Peninsular Malaysia's most important conservation area. Estimated densities ( ) of adult tigers ranged from 1.10 ± 0.52 to 1.98 ± 0.54 tigers/100 km2 (X2=1.56, df=2, P=0.46) with the overall mean of 1.66 ± 0.21 tigers/100 km2. The tiger population in the 4343-km2 park was estimated to be 68 (95% CI: 52–84) adult tigers. Prey biomass estimates ranged from 266 to 426 kg/km2, and wild boar were the most important potential prey species in terms of abundance, biomass, and occupancy, followed by muntjac. Both tigers and leopards were more diurnal than nocturnal, which corresponded with the activity patterns of wild boar and muntjac. No evidence of poaching of large mammals was found in the 600-km2 study sites and overall human impacts on the tiger–prey community appear to be minimal, but in the long run its viability needs to be evaluated in a greater landscape context.
Article
We investigated introgression in a mixed weedy population of oilseed rape (Brassica napus) and its relative B. rapa using species-specific AFLP-markers. The population was situated in a field relayed from conventional to organic cultivation 11 years ago. One-hundred-and-two B. napus or B. rapa-like plants were collected in a 3 m^2 plot. Of these, one was a first generation hybrid (F_1) and nearly half (44 plants) were introgressed, having both B. napus and B. rapa specific markers. The remaining plants apparently corresponded to pure species, with 50 having only B. rapa- and seven having only B. napus-specific markers. We compared the number of markers in the plants from the weedy population with the numbers in controlled backcross generations (BC_1 and BC_2). The marker distribution in the weedy population resembled the distribution in the second backcross generation most closely. Together with the cultivation history of the field, this suggests that the introgression process in the weedy mixed population has been in progress for some time. This study is the first to show introgression between B. napus and B. rapa under natural conditions.
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Genomic paw prints in the DNA of the world’s wild cats have clarified the cat family tree and uncovered several remarkable migrations in their past
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Wildlife managers require status and distribution information for informed decisions. Recognizing the tiger's globally threatened status and potential as an umbrella species for protection of forested landscapes, camera trap surveys for tigers and other large mammals have been conducted since 1997 in Peninsular Malaysia with the aim of assessing the population status of tigers in the Peninsula. Results from surveys at nine sites between December 1997 and December 1999 are reported here. Tigers were confirmed from six sites in the Main Range and Greater Taman Negara landscape, with multiple locations inside putative priority tiger areas. Although the data were collected 8 years ago, they are supplemented with more recent information, including tiger-human conflict investigations during 2000–2005 that indicate tiger persistence at these sites. Tiger density estimates were 0.51–1.95 tigers per 100 km 2 . With results from other surveys, this suggests a national population of up to several hundred tigers. A thorough survey, with sufficient resources, should be carried out in the future to derive a more reliable tiger population estimate for Malaysia. Key threats are habitat loss and fragmentation, hunting of prey, commercial trade in tiger parts, and harassment and displacement. Recommendations for the recovery of tigers in Peninsu-lar Malaysia are provided.
Article
The black form of the leopard is shown to be inherited as an autosomal monogenic recessive to the spotted wild type. Pairings of black animals inter se have a significantly smaller litter size than other possible pairings. Two reasons are offered for this: (1) that black animals may be slightly more inbred than the spotted or (2) black animals may be more prone to pre-natal loss than the spotted. The significance of the latter possibility is briefly discussed in relation to the persistence of the spotted/black polymorphism for areas of South-east Asia.
Article
The effect of the Toba super-eruption at ∼74 kyr on the mammals of Southeast Asia is examined. Although few Late Pleistocene sites from Southeast Asia have been described, an analysis of those which pre- and post-date Toba reveals relatively few species became extinct following the eruption. It is suggested that species survived in refugia immediately following the eruption, and that they repopulated vast areas following a probable short period (i.e. decades to century) of environmental devastation. This study suggests that mammals are more robust at coping with catastrophic events than previously acknowledged, and questions the perceived human monopoly in overcoming ecological adversity.
Article
Thesis (M.S.)--University of Minnesota, 1996. Includes bibliographical references (leaves 54-63).
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There is abundant geographic variation in both morphology and gene frequency in most species. The extent of geographic variation results from a balance of forces tending to produce local genetic differentiation and forces tending to produce genetic homogeneity. Mutation, genetic drift due to finite population size, and natural selection favoring adaptations to local environmental conditions will all lead to the genetic differentiation of local populations, and the movement of gametes, individuals, and even entire populations--collectively called gene flow--will oppose that differentiation. Gene flow may either constrain evolution by preventing adaptation to local conditions or promote evolution by spreading new genes and combinations of genes throughout a species' range. Several methods are available for estimating the amount of gene flow. Direct methods monitor ongoing gene flow, and indirect methods use spatial distributions of gene frequencies to infer past gene flow. Applications of these methods show that species differ widely in the gene flow that they experience. Of particular interest are those species for which direct methods indicate little current gene flow but indirect methods indicate much higher levels of gene flow in the recent past. Such species probably have undergone large-scale demographic changes relatively frequently.
Article
Melanistic coat coloration occurs as a common polymorphism in 11 of 37 felid species and reaches high population frequency in some cases but never achieves complete fixation. To investigate the genetic basis, adaptive significance, and evolutionary history of melanistic variants in the Felidae, we mapped, cloned, and sequenced the cat homologs of two putative candidate genes for melanism (ASIP [agouti] and MC1R) and identified three independent deletions associated with dark coloration in three different felid species. Association and transmission analyses revealed that a 2 bp deletion in the ASIP gene specifies black coloration in domestic cats, and two different "in-frame" deletions in the MC1R gene are implicated in melanism in jaguars and jaguarundis. Melanistic individuals from five other felid species did not carry any of these mutations, implying that there are at least four independent genetic origins for melanism in the cat family. The inferred multiple origins and independent historical elevation in population frequency of felid melanistic mutations suggest the occurrence of adaptive evolution of this visible phenotype in a group of related free-ranging species.
Aborigines in Taman Negara: the impact of a hunters, gatherers, community on a low-land rainforest ecosystem
  • Van
  • W E M Schot
Van der Schot, W.E.M. (1990). Aborigines in Taman Negara: the impact of a hunters, gatherers, community on a low-land rainforest ecosystem. J. Wildl. Parks (Malaysia) 10, 172–183
c ?2010 The Authors Journal of Zoology c ?2010 The Zoological Society of London 205 Melanistic leopards of the Malay Peninsula K Udjung Kulon, the land of the last Javan rhinoceros A comparison of amphibian communities through time and from place to place in Bornean forests
  • Kawanishi
Journal of Zoology 282 (2010) 201–206 c ?2010 The Authors. Journal of Zoology c ?2010 The Zoological Society of London 205 Melanistic leopards of the Malay Peninsula K. Kawanishi et al. rHoogerwerf, A. (1970). Udjung Kulon, the land of the last Javan rhinoceros. Leiden: E. J. Brill. Inger, R.F. &Voris, H.K. (1993). A comparison of amphibian communities through time and from place to place in Bornean forests. J. Trop. Ecol. 9, 409–433
The ecology and conservation of tigers, other large carnivores, and their prey in Kuiburi National Park, Thai-land
  • R Steinmetz
  • W Chutipong
  • N Seuaturien
  • B Poonnil
Steinmetz, R., Chutipong, W., Seuaturien, N. & Poonnil, B. (2007). The ecology and conservation of tigers, other large carnivores, and their prey in Kuiburi National Park, Thai-land. Bangkok: WWF Thailand and Department of Na-tional Parks, Wildlife, and Plant Conservation.
The big cats and their fossil relatives Aborigines in Taman Negara: the impact of a hunters, gatherers, community on a low-land rainforest ecosystem
  • A Turner
  • Van
  • W E M Schot
Turner, A. (1997). The big cats and their fossil relatives. New York: Columbia University Press. Van der Schot, W.E.M. (1990). Aborigines in Taman Negara: the impact of a hunters, gatherers, community on a low-land rainforest ecosystem. J. Wildl. Parks (Malaysia) 10, 172–183.
Rarity and possible new records of Sumatran rhinoceros in Taman Negara
  • K Kawanishi
  • M Sunquist
  • A M Sahak
Kawanishi, K., Sunquist, M. & Sahak, A.M. (2002). Rarity and possible new records of Sumatran rhinoceros in Taman Negara. J. Wildl. Parks (Malaysia) 20, 125–128.
Ecology and conservation of tigers and their prey in Kuiburi National Park
  • R Steinmetz
  • W Chutipong
  • N Seuaturien
  • B Poonnil
Steinmetz, R., Chutipong, W., Seuaturien, N. & Poonnil, B. (2009). Ecology and conservation of tigers and their prey in Kuiburi National Park, Thailand. Bangkok: WWF Thai-land and Department of National Parks.
Udjung Kulon, the land of the last Javan rhinoceros
  • A Hoogerwerf
Hoogerwerf, A. (1970). Udjung Kulon, the land of the last Javan rhinoceros. Leiden: E. J. Brill.
The wild mammals of Malaya (Peninsular Malaysia) and Singapore Mammal diversity and conservation in a secondary forest of peninsular Malaysia
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Medway, L. (1978). The wild mammals of Malaya (Peninsular Malaysia) and Singapore, 2nd edn. Petaling Jaya: Oxford University Press. Mohd Azlan, J. (2006). Mammal diversity and conservation in a secondary forest of peninsular Malaysia. Biodivers. Conserv. 15, 1013–1025.
Distributions of large fauna with respect to the edge of a Thailand protected area
  • A J Lynam
Lynam, A.J. (1996). Distributions of large fauna with respect to the edge of a Thailand protected area. Unpublished report to Wildlife Conservation Society, New York.
An account of the wild tribes inhabiting the Malayan Peninsula, and a few neighboring islands. Paris: Imperial Printing Office
  • P E L Favre
Favre, P.E.L. (1865). An account of the wild tribes inhabiting the Malayan Peninsula, and a few neighboring islands. Paris: Imperial Printing Office.
A preliminary survey of tiger and other large mammals at Kaeng Krachan national park, Petchburi province, Thailand
  • Ngoprasert D.
Ngoprasert, D. & Lynam, A.J. (2002). A preliminary survey of tiger and other large mammals at Kaeng Krachan national park, Petchburi province, Thailand. J. Wildl. Thailand 10, 33–38 (in Thai).
Tiger Monitoring Study in Gunung Basor Forest Reserve, Molecular genetics and evolution of melanism in the cat family
  • M R Darmaraj
  • Jeli
  • Kelantan
  • Wwf-Malaysia
  • Jaya
  • Malaysia
  • E Eizirik
  • N Yuhki
  • W E Johnson
  • M Menotti-Raymond
  • S S Hannah
  • J Brien
Darmaraj, M.R. (2007). Tiger Monitoring Study in Gunung Basor Forest Reserve, Jeli, Kelantan. WWF-Malaysia, Pe-taling Jaya, Malaysia. Eizirik, E., Yuhki, N., Johnson, W.E., Menotti-Raymond, M., Hannah, S.S. & O'Brien, J. (2003). Molecular genetics and evolution of melanism in the cat family. Current Biol. 13, 448–453.
The breeding of spotted and black leopards
  • Robinson R.
Robinson, R. (1969). The breeding of spotted and black leopards. J. Bombay Nat. Hist. Soc. 66, 423–429.
Tiger Monitoring Study in Gunung
  • M R Darmaraj
Darmaraj, M.R. (2007). Tiger Monitoring Study in Gunung Basor Forest Reserve, Jeli, Kelantan. WWF-Malaysia, Petaling Jaya, Malaysia.
  • H P Goudriaan
Goudriaan, H.P. (1948). Panters. Indische Jager 2, 77–90.
  • Boomgaard
Using remote photography in wildlife ecology
  • Cutler T.L.
Black panthers – an inquiry
  • Pocock R.I.