Density and microhabitat use of Bengal slow loris in primary forest and non-native plantation forest

The extent of planted forests has greatly increased in the tropics, but their conservation value while assumed to be low, is largely unknown. We compared the density and microhabitat selection of a nocturnal arboreal primate, the Bengal slow loris (Nycticebus bengalensis), in mostly undisturbed, evergreen tropical forest to those in 15-18 year old Acacia/Leucaena plantations with significant secondary regrowth, and <15 year old plantations with little regrowth. Based on estimates derived from distance sampling, loris densities in older plantations were nearly identical to primary forest (4.26 vs. 4.00 lorises per square kilometer), although encounter rates were three times higher in the older plantations probably owing to the lower detection probability in the more complex vegetation of the primary forest. The mean density estimate for the younger plantation was one-third of the above habitats (1.27 lorises per square kilometer), although not statistically different. Lorises tended to use larger diameter and taller trees, with a greater crown depth than randomly sampled trees, and tended to avoid habitats with sparsely crowned trees. The older plantations had trees with lower basal area and shorter stems than the primary forest; however, the older plantations contained higher densities of Bauhinia lianas, a commonly eaten food source and did not contain the red giant flying squirrel (Petaurista petaurista), a potential competitor. Although it is unknown whether the Bengal slow loris would persist without the presence of primary forest in the landscape, we suggest that older plantations have conservation value for at least selected species and as such, could be better managed to increase this value.     

Exudativory in the Bengal slow loris (Nycticebus bengalensis) in Trishna Wildlife Sanctuary,Tripura, northeast India

In this study we estimated the extent of exudativory in Nycticebus bengalensis and examined whether exudates can be considered as fallback foods. This study was carried out in Trishna Wildlife Sanctuary, northeastern India, in winter (December–February) and summer (March and April). We estimated time–activity budget using instantaneous sampling and used continuous focal animal sampling to record all instances and durations of feeding, over a total of 177 hr. Feeding accounted for 22.3±2.2% of the activity budget, with no seasonal difference. Bengal slow lorises fed on exudates, nectar, fruit, bark, invertebrates and avian eggs. In addition to scraping they also obtained exudates by gouging holes into the bark of trees. In winter, lorises almost exclusively fed on exudates (94.3% of winter feeding time). In summer, exudates (67.3%) and nectar from one species (22.3%) dominated the diet. This study identifies the Bengal slow loris as the most exudativorous loris. Exudates rather than being a staple fallback food, seem to be a preferred, patchily distributed and common food in the diet of the Bengal slow loris. Exudativory in this species is characterized by high selectivity among species and seasonal variation, which may be related to variations in productivity of exudates and their chemical composition. An understanding of these factors is necessary for predicting the response of this species to human disturbance such as logging. This study also underscores the importance of protecting some of the common species such as Terminalia belerica on which the loris feeds during periods of scarcity. Am. J. Primatol. 72:113–121, 2010. © 2009 Wiley‐Liss, Inc.     

Distribution and Population Densities of Diurnal Primates in the Karst Forests of Phong Nha – Ke Bang National Park,Quang Binh Province,Central Vietnam

We studied the status and distribution of the diurnal primates in the Phong Nha – Ke Bang National Park (PNKB NP) from April to August 2007. In the past, researchers reported 9 primate species and subspecies for the karst forests of PNKB NP, which constitutes the most important protected area for the endangered Hatinh langur (Trachypithecus hatinhensis) in Vietnam. All 9 primate taxa are threatened due to hunting pressure and habitat loss. We applied line transect sampling in 4 areas. During a total of 117 transect inspections along 12 different transect routes, we confirmed 5 primate taxa and the analysis of population densities resulted in 2143 (±467) Hatinh langurs, 1316 (±871) red-shanked douc langurs (Pygathrix nemaeus), 930 (±489) stump-tailed macaques (Macaca arctoides), 986 (±883) eastern Assamese macaques (M. a. assamensis), and 18 (±18) southern white-cheeked crested gibbons (Nomascus siki) in the whole PNKB NP, which covers an area of ca. 85,000 ha. We could not detect the 2 nocturnal lorises, Bengal slow lorises (Nycticebus bengalensis) and pygmy slow lorises (N. pygmaeus), as well as rhesus macaques (Macaca mulatta) and northern pig-tail macaques (M. leonina). The distribution of the primates predominantly depended on human impact. We could not recognize a correlation between habitat constitution and abundance of primates. The population density estimates showed a much higher density of the Hatinh langur than previously assumed. Thus the importance of the PNKB NP for the conservation of this endangered langur increased significantly.     

Mother-infant interactions in slow lorises (Nycticebus bengalensis) and pygmy lorises (Nycticebus pygmaeus)

The study had three purposes: (1) to obtain information about mother-infant interactions in a rarely studied nocturnal prosimian, the pygmy loris (Nycticebus pygmaeus); (2) to compare pygmy lorises with a closely related and better-studied nocturnal prosimian, the Bengal slow loris (Nycticebus bengalensis); and (3) to determine how the presence of a second offspring affected mother-infant interactions in pygmy lorises. Three Bengal slow loris mothers and 3 pygmy loris mothers served as subjects, along with their 10 offspring (4 Bengal slow loris singletons, 2 pygmy loris singletons and 2 sets of pygmy loris twins). Observations were carried out in a zoo research facility for the first 24 weeks of the infants' lives. Although the two species differ in size and reproductive patterns, mother-infant interactions were similar. The primary modes of infant and adult contact were ventral and passive contact, respectively. Mothers parked their infants from the first week, and infants followed from the second week. Mothers displayed little protection or rejection, and there was little aggression. Infants solicited play and social grooming from their mothers. Pygmy loris mothers engaged in social grooming and play with their infants more frequently and for longer periods if the infant was a singleton rather than a twin.     

Artificial canopy bridges improve connectivity in fragmented landscapes: The case of Javan slow lorises in an agroforest environment

Canopy bridges are increasingly used to reduce fragmentation in tropical habitats yet monitoring of their impact on the behavior of primates remains limited. The Javan slow loris (Nycticebus javanicus) is endemic to Java, Indonesia, where the species most often occurs in human-dominated, highly patchy landscapes. Slow lorises cannot leap, are highly arboreally adapted, and are vulnerable on the ground. To increase arboreal connectivity, as part of a long-term conservation project in Cipaganti, West Java, we built and monitored seven slow lorises bridges of two types—waterline or rubber—and monitored their use by seven adult individuals from 2016 to 2017. Motion triggered camera traps collected data for 195 ± standard deviation (SD) 85 days on each bridge. We collected 341.76 hr (179.67 hr before and 162.09 hr after the installation of bridges) of behavioral and home range data via instantaneous sampling every 5 min, and terrestrial behavior (distance and duration of time spent on the ground) via all occurrences sampling. We found that slow lorises used bridges on average 12.9 ± SD 9.7 days after their installment mainly for traveling. Slow lorises showed a trend toward an increase in their home range size (2.57 ha before, 4.11 ha after; p = 0.063) and reduced ground use (5.98 s/hr before, 0.43 s/hr; p = 0.063) after implementation of bridges. Although the number of feeding trees did not change, new feeding trees were included in the home range, and the proportion of data points spent traveling and exploring significantly decreased (p = 0.018). Waterline bridges serve a purpose to irrigate the crops of local farmers who thus help to maintain the bridges, and also ascribe value to the presence of slow lorises. Other endemic mammal species also used the bridges. We advocate the use and monitoring of artificial canopy bridges as an important supplement for habitat connectivity in conservation interventions.     

A Survey of Husbandry Practices for Lorisid Primates in North American Zoos and Related Facilities

Zoos and related facilities in North America currently manage five species in the primate family Lorisidae: the greater (Nycticebus coucang), Bengal (N. bengalensis) and pygmy (N. pygmaeus) slow lorises, red slender loris (Loris tardigradus), and potto (Perodicticus potto). We used an online survey to describe institutional housing and husbandry practices for these species and assess the extent to which practices are consistent with established guidelines. Our results show that most captive lorisids are housed solitarily or in pairs. Most individuals occupy a single exhibit space in a building dedicated to nocturnal animals. Facilities are commonly meeting recommendations for abiotic exhibit design and are providing animals with an enriched environment. However, pottos and slender lorises currently occupy exhibit spaces smaller than the recommended minimum, and the impact of cleaning protocols on olfactory communication should be critically evaluated. Few facilities are taking advantage of the benefits of positive reinforcement training for promoting animal welfare. Research is greatly needed on the effects of exhibit lighting on behavior, health, and reproduction; and to determine how best to manage the social needs of lorisids with naturally dispersed social structures. Although captive populations of slender lorises, pottos, and slow lorises are declining, we suggest that improved husbandry knowledge has the potential to positively influence population sustainability and to enhance future efforts to manage the growing pygmy loris population. Zoo Biol. 32:88‐100, 2013. © 2012 Wiley Periodicals, Inc.     

Conservation implications of low encounter rates of five nocturnal primate species (<Emphasis Type="Italic">Nycticebus </Emphasis>spp.) in Asia

Five species of slow lorises were once considered to comprise a single strongly polymorphic species, Nycticebus coucang, ranging throughout South and Southeast Asia. The cryptic nature of these nocturnal primates has led to a lack of understanding of their distribution patterns and abundance. In short surveys, often few if any lorises are detected, meaning that the few available density estimates are from long-term studies. Based on new research in Sebangau National Park, Borneo, and compilation of survey data from other areas, we provide the first comparative abundance estimates for all five slow loris species: N. coucang occurred in significantly higher abundances (median encounter rate 0.80/km: n = 15), than N. bengalensis (0.26/km; n = 12), or N. javanicus (0.11/km: n = 2), N. menagensis (0.02/km: n = 3), and N. pygmaeus (0.13/km: n = 4). Abundance estimates in Sebangau (0.19/km) did not increase with increasing survey effort, but for all species and studies combined, study duration was positively correlated with abundance estimates. We did not find a relation between abundance and body mass, nor between abundance and latitude. Long-term studies are more likely to be conducted at sites where the species of interest is particularly plentiful. The data suggest that slow lorises occur at low abundances throughout much of their range, and some in larger social groups than previously assumed. We recommend taking into account the species’ heterogeneous distribution (potentially requiring larger survey effort), their social structure, the use of red lights as opposed to white lights whilst surveying, and to make use of their vocalisations when surveying slow lorises.     

Phylogeny of the slow lorises (genusNycticebus): An approach using mitochondrial DNA restriction enzyme analysis

Mitochondrial DNA polymorphisms in 15 specimens of three species of slow lorises-Nycticebus coucang, N. intermedius, andN. pygmaeus-were analyzed in order to study the evolutionary relationships among the species. Eight restriction types were observed in the samples. Phylogenetic trees constructed on the basis of genetic distances showed that the slow lorises sort into two clusters: four types ofN. coucang and three types ofN. intermedius plus one type ofN. pygmaeus. Our results suggest that there are two valid species in the genusNycticebus-N. coucang, andN. pygmaeus-and thatN. intermedius should be included withinN. pygmaeus. Divergence between the two species may have begun 2.7 Ma (million years ago). Evolution of gross morphology, chromosomes, and mitochondrial DNA in the slow lorises appears to be concordant.     

Distribution and Conservation of Slender Lorises (Loris tardigradus lydekkerianus) in Southern Andhra Pradesh,South India

Slender lorises live in forests of southern India and Sri Lanka. Little is known about their distribution patterns and relative densities. We report the results of a survey conducted in 6 forest divisions in the southern parts of the state of Andhra Pradesh, South India. Relatively high densities of lorises occurred in mixed deciduous forests and in adjoining farm lands interspersed with trees. Three distinct populations inhabit the study area. We recommend conservation measures for Loris tardigradus.     

Mitochondrial Control Region and Population Genetic Patterns of Nycticebus bengalensis and N. pygmaeus

Bengal slow lorises (Nycticebus bengalensis) and pygmy slow lorises (Nycticebus pygmaeus) are nocturnal which creates difficulties to study them in the field. There is a scarcity of data on them and their population genetics are poorly understood. We sequenced and analyzed a partial fragment in the first hypervariable region of the mitochondrial control region or D-loop HVRI of 21 Nycticebus bengalensis and 119 N. pygmaeus from the boundary between China and Vietnam where they are sympatric. Though the sample size for Nycticebus pygmaeus is much larger, the polymorphism level is much lower than that of N. bengalensis, possibly because of (1) external gene flow from other habitats of N. bengalensis, (2) gene ingression from Sunda slow lorises (N. coucang coucang) to N. bengalensis, (3) a skewed birth sex ratio in N. pygmaeus, and (4) a possible low survival rate of infant N. pygmaeus. Based on genetic comparisons to Nycticebus bengalensis, we propose that N. pygmaeus in southern China and northern Vietnam might have migrated from middle or southern Vietnam recently. Deng Pan and Jing-Hua Chen made equal contributions to the work.     

Coexistence between Javan Slow Lorises (<Emphasis Type="Italic">Nycticebus javanicus</Emphasis>) and Humans in a Dynamic Agroforestry Landscape in West Java,Indonesia

In a world increasingly dominated by human demand for agricultural products, we need to understand wildlife’s ability to survive in agricultural environments. We studied the interaction between humans and Javan slow lorises (Nycticebus javanicus) in Cipaganti, Java, Indonesia. After its introduction in 2013, chayote (Sechium edule), a gourd grown on bamboo lattice frames, became an important cash crop. To evaluate people’s use of this crop and to measure the effect of this increase on slow loris behavior, home ranges, and sleep sites, we conducted interviews with local farmers and analysed the above variables in relation to chayote expansion between 2011 and 2015. Interviews with farmers in 2011, 2013, and 2015 confirm the importance of chayote and of bamboo and slow lorises in their agricultural practices. In 2015 chayote frames covered 12% of land in Cipaganti, occupying 4% of slow loris home ranges, which marginally yet insignificantly increased in size with the increase in chayote. Slow lorises are arboreal and the bamboo frames increased connectivity within their ranges. Of the sleep sites we monitored from 2013 to 2016, 24 had disappeared, and 201 continued to be used by the slow lorises and processed by local people. The fast growth rate of bamboo, and the recognition of the value of bamboo by farmers, allow persistence of slow loris sleep sites. Overall introduction of chayote did not result in conflict between farmers and slow lorises, and once constructed the chayote bamboo frames proved to be beneficial for slow lorises.     

Slow Lorises (<Emphasis Type="Italic">Nycticebus</Emphasis> spp.) Really Are Slow: a Study of Food Passage Rates

The characteristics of food ingested by a primate affect its assimilation of energy by modulating food passage rate. In general, digestive time increases in folivorous primates and decreases in frugivorous primates when they are fed higher fiber diets but this relationship is understudied in exudativorous primates. We compared the food passage rate of five slow loris species. We studied 34 wild-caught slow lorises (15 Nycticebus coucang, 15 N. javanicus, and 4 N. menegensis) in an Indonesian rescue center and four captive-born slow lorises (2 N. bengalensis and 2 N. pygmaeus) in a UK institution. We fed the Indonesian subjects two different diets: a captive-type diet comprising fruits, vegetables, and insects and a wild-type diet formulated to be similar in nutrients to that consumed by slow lorises in the wild, consisting of gum, insects, vegetables, and nectar. We fed the UK subjects a diet of gum, vegetables, insects, and hard-boiled eggs. We formulated this diet to mimic the wild diet, with notably higher fiber fractions and lower soluble sugars than the previous diet. We measured two variables: the transit time (TT) and the mean retention time (MRT). We mixed 1 tsp. of glitter in bananas or gum as our markers and fed them to the slow lorises immediately before their main diet. We noted the date and time of feeding and of appearances of the marker in feces. We weighed food given and left over for each individual to calculate ingested foods and nutrients. We found that TTs were not affected by diet treatment but MRTs were significantly longer for all species fed the wild-type diet. Our results show that Nycticebus spp. have long MRTs for their body weight, and N. pygmdaeus may have the slowest MRT of all primates in relation to body mass. The digestive flexibility of exudativorous primates should allow them to maximize fermentation opportunities when they ingest more (appropriate) fiber by increasing the amount of time the fiber substrate stays in the large intestine. Exudativorous primates appear to have plastic digestive strategies that may be an adaptation to cope with relatively nutrient-poor staple food sources such as gum. The provision of gum in a captive setting may therefore provide benefits for gut health in slow lorises.     

Checkerboard Patterns,Interspecific Competition,and Extinction: Lessons from Distribution Patterns of Tarsiers (<Emphasis Type="Italic">Tarsius</Emphasis>) and Slow Lorises (<Emphasis Type="Italic">Nycticebus</Emphasis>) in Insular Southeast Asia

Tarsiers (Tarsius) and slow lorises (Nycticebus) are the only extant nocturnal primates occurring in Southeast Asia. Harcourt (1999) hypothesized that in insular Southeast Asia, slow lorises and tarsiers showed a checkerboard distribution on 12 small (<12,000 km²) islands, i.e., only one or the other occurs, and attributed this to extreme levels of competition between these 2 largely faunivorous primates. Further, he predicted slow lorises were able to persist on smaller islands than tarsiers. We re-evaluated these findings using an expanded dataset including 49 islands where tarsiers or slow lorises occur. Tarsiers and slow lorises live on islands of similar size (median size of ca. 300–900 km²), and both taxa inhabit an equal proportion of small, medium, and large islands. On small islands within their area of sympatry tarsiers occur on 1 island, slow lorises on 8, both genera on 3, and we can assume they have become extinct from 11 small islands since the Last Glacial Maximum. Sizes of islands where tarsiers or slow lorises have become extinct do not differ from islands where they are still extant. We show that slow lorises occur on more islands in insular Southeast Asia than perhaps previously assumed, but these islands are not smaller on average than islands where tarsiers occur. A checkerboard distribution between these taxa is not evident. More studies are needed at the macroecological level to assess the importance of biogeographic history in explaining their present-day distribution patterns.     

Extreme primates: Ecology and evolution of Asian lorises

Asia's slow and slender lorises (Nycticebus and Loris) are among nature's most extreme primates. Until recently, it was not understood why lorises have such huge forward‐facing eyes, strange steady climbing locomotion, tiny dependent babies, and a bite that potentially can kill a human! Indeed, early studies described them as slow, solitary, and boring. Twenty years of field research now indicate that lorises are among the most intriguing mammal species.     

Welfare impacts of the illegal wildlife trade in a cohort of confiscated greater slow lorises,Nycticebus coucang

Illegal harvesting and trade are major forces behind population declines of wild slow lorises (genus Nycticebus). The impacts of the wildlife trade on individual slow lorises have not been as well described. In this article, we describe quantitatively the consequences of the wildlife trade for 77 greater slow lorises, N. coucang, who were confiscated en masse and brought to Cikananga Wildlife Center in Indonesia. Medical records indicated that in total, 28.6% of the slow lorises died within the first 6 months, mostly due to traumatic injury, and all the infants died. The greatest sources of morbidity were external wounds (33.1% of 166 total medical events) and dental problems (19.3%). Of the surviving individuals, 25.4% displayed abnormal behavior. Behavioral observations indicated that healthy adults (n = 3) spent 48.2% of their active period performing stereotypies. These data illustrate the physical and behavioral impacts of the illegal wildlife trade on the welfare of slow lorises. We suggest that sharing these individual stories may help generate empathy and educate the public about the impacts of the exotic companion-animal (pet) trade on nonhuman animal welfare.     

Aposematic signaling and seasonal variation in dorsal pelage in a venomous mammal

In mammals, colouration patterns are often related to concealment, intraspecific communication, including aposematic signals, and physiological adaptations. Slow lorises (Nycticebus spp.) are arboreal primates native to Southeast Asia that display stark colour contrast, are highly territorial, regularly enter torpor, and are notably one of only seven mammal taxa that possess venom. All slow loris species display a contrasting stripe that runs cranial‐caudally along the median sagittal plane of the dorsum. We examine whether these dorsal markings facilitate background matching, seasonal adaptations, and intraspecific signaling. We analyzed 195 images of the dorsal region of 60 Javan slow loris individuals (Nycticebus javanicus) from Java, Indonesia. We extracted greyscale RGB values from dorsal pelage using ImageJ software and calculated contrast ratios between dorsal stripe and adjacent pelage in eight regions. We assessed through generalized linear mixed models if the contrast ratio varied with sex, age, and seasonality. We also examined whether higher contrast was related to more aggressive behavior or increased terrestrial movement. We found that the dorsal stripe of N. javanicus changed seasonally, being longer and more contrasting in the wet season, during which time lorises significantly increased their ground use. Stripes were most contrasting in younger individuals of dispersal age that were also the most aggressive during capture. The dorsal stripe became less contrasting as a loris aged. A longer stripe when ground use is more frequent can be related to disruptive colouration. A darker anterior region by younger lorises with less fighting experience may allow them to appear larger and fiercer. We provide evidence that the dorsum of a cryptic species can have multimodal signals related to concealment, intraspecific communication, and physiological adaptations.     

Distribution Patterns of Slender Loris Subspecies (<Emphasis Type="Italic">Loris lydekkerianus</Emphasis>) in Kerala,Southern India

Gray slender lorises (Loris lydekkerianus) are 1 of 2 nocturnal primate genera occurring in India. Although the distribution and abundance of the species is known for some parts of southern India, the relative distribution of the 2 subspecies, Loris lydekkerianus lydekkerianus and L. l. malabaricus, and their comparative densities and extent of overlap between their distributions remains largely unknown. We investigated the distribution pattern and relative density of slender lorises in the Western Ghats mountain range in the state of Kerala in southern India. We surveyed 36 forest ranges in 17 forest divisions of northern and central Kerala from February to November 2009 for the presence of lorises. We sighted slender lorises in 22 forest ranges in a variety of vegetation types, and the relative abundance of the species ranged from 0.02 to 1.44 lorises/km. Our results confirm that both loris subspecies are present in Kerala: Mysore slender lorises have a narrow patchy distribution on the eastern edges of the Western Ghats mountain range, and Malabar slender lorises have a fairly contiguous distribution on the western slopes of the Western Ghats. We recommend more detailed surveys in southern Kerala to identify the distribution patterns of the subspecies in these areas.     

Tickled to Death: Analysing Public Perceptions of ‘Cute’ Videos of Threatened Species (Slow Lorises – Nycticebus spp.) on Web 2.0 Sites

Background

The internet is gaining importance in global wildlife trade and changing perceptions of threatened species. There is little data available to examine the impact that popular Web 2.0 sites play on public perceptions of threatened species. YouTube videos portraying wildlife allow us to quantify these perceptions.

Methodology/Principal Findings

Focussing on a group of threatened and globally protected primates, slow lorises, we quantify public attitudes towards wildlife conservation by analysing 12,411 comments and associated data posted on a viral YouTube video ‘tickling slow loris’ over a 33-months period. In the initial months a quarter of commentators indicated wanting a loris as a pet, but as facts about their conservation and ecology became more prevalent this dropped significantly. Endorsements, where people were directed to the site by celebrities, resulted mostly in numerous neutral responses with few links to conservation or awareness. Two conservation-related events, linked to Wikipedia and the airing of a television documentary, led to an increase in awareness, and ultimately to the removal of the analysed video.

Conclusions/Significance

Slow loris videos that have gone viral have introduced these primates to a large cross-section of society that would not normally come into contact with them. Analyses of webometric data posted on the internet allow us quickly to gauge societal sentiments. We showed a clear temporal change in some views expressed but without an apparent increase in knowledge about the conservation plight of the species, or the illegal nature of slow loris trade. Celebrity endorsement of videos showing protected wildlife increases visits to such sites, but does not educate about conservation issues. The strong desire of commentators to express their want for one as a pet demonstrates the need for Web 2.0 sites to provide a mechanism via which illegal animal material can be identified and policed.     

Studies on the chromosomes of genusNycticebus

The karyotypes of three species (N. coucang, N. intermedius, andN. pygmaeus) of genusNycticebus, collected from the southern Yunnan of China, have been studied. All individuals from three species possess 2n=50 chromosomes, and all chromosomes in their complement are biarm chromosomes. The karyotype of slow loris (N. coucang) is characterized by having a secondary constriction and Ag-NORs in the short arms of pair No. 1. The G-banding patterns of three species are very similar. Three species are found to have multiple Ag-NORs. InN. coucang, NORs were observed on five pairs (Nos. 1, 6, 9, 15, and 23) and inN. intermedius andN. pygmaeus, NORs were found on four pairs (Nos. 6, 9, 15, and 20). This finding indicates that slow lorises, as primitive primates, also have multiple NOR-bearing chromosomes. Finally, the classification of genusNycticebus by karyotype analysis is discussed, and our results suggest that there are at least two valid species, namely:N. coucang andN. pygmaeus.     

中国兰科一新记录种——毛瑞线柱兰

报道了在海南热带雨林国家公园毛瑞管理分局发现的兰科线柱兰属一中国新记录种——毛瑞线柱兰(Zeuxine glandulosa King&Pantl).该种原记录分布于印度阿萨姆邦、锡金邦,北孟加拉和泰国.本文对该种形态及生境进行了详尽描述,并提供高清解剖照及线描图.凭证标本保存于海南大学林学院教学标本馆(HUFB).