Staff Profiles

Modules taught

Undergraduate

• Undergraduate: Methods and Analysis in Biological Anthropology
• Undergraduate: Introduction to Biological Anthropology

Postgraduate

• Postgraduate: Primate Conservation - Conservation Education
• Postgraduate: Independent study in lemurs and nocturnal primates
• Postgraduate: MSc Primate Conservation Final Project

Supervision

Postdoctoral research Fellows

• Dr Marie Sigaud: understanding resource selection by Javan slow lorises in an ever-changing agroforest in West Java

Research students

• Francis Cabana, Wild Bites:  Using Nutritional Ecology (Native Ingredients and Chemical Composition) of the Javan Slow Loris  to Improve Captive Feeding Husbandry
• Stephanie Poindexter: Cognition and Conservation: measuring spatial cognition in slow lorises and its application to reintroduction practices in Vietnam, Thailand and Indonesia
• Katie Reinhardt: Energetics in a changing climate: implications for behavioural and evolutionary adaptations of the Javan slow loris
• Averee Luhrs: Perodicticinae and primate origins: investigating the niche of the pottos and angwantibos in the context of evolutionary theory with implications for modern-day management
• Miguel de Guinea: Cognitive maps in black howler monkeys at Palenque National Park, Mexico

I focus on the behaviour, ecology, taxonomy and conservation of nocturnal mammals. I work mainly in Asia, where I have conducted research in 12 countries. I am a long-standing member of the Nocturnal Primate Research Group and Director of the Little Fireface Project, a charity and research project that works to conserve understudied obscure nocturnal mammals. The main countries where I have conducted my work are India (slow and slender lorises); Sri Lanka (langurs, macaques, slender lorises, civets, small cats); Cambodia (pygmy slow lorises) and Indonesia, where I now run a long-term project on the Critically Endangered slow loris. Topics of my work have included: evolution of venom in mammals, thermoregulation in slow lorises, evolution of exudativory in primates, and using morphology to distinguish species. 

Research interests

• Conservation
• Behavioural Ecology
• Speciation
• Fragmentation
• Nocturnal mammals and colobines

Books

• , Ed., Evolution, ecology and conservation of lorises and pottos, (2020)
  ISBN: 9781108429023 eISBN: 9781108676526
  Abstract

  Furry and wide-eyed, lorises and pottos are small, nocturnal primates inhabiting African, Asian and Southeast Asian tropical and subtropical forests. Their likeable appearance, combined with their unusual adaptations - from a marked reduction of the tail to their mostly slow, deliberate locomotion, powerful grasping and, in some species, a venomous bite - has led to a significant rise in research interest in the family Lorisidae over the last decade. Furthermore, lorises in particular have featured frequently in international media largely due to illegal trade, for example as pets. This is the first volume to present a full picture of the breadth of research being undertaken on lorisids to aid future studies as well as conservation efforts. Focusing on five key topics: evolutionary biology, ecomorphology, behavioural ecology, captive management and conservation, this book is a vital read for graduate students and researchers in primatology, biological anthropology, evolutionary biology, animal behaviour and conservation.
  

  Website 

Journal articles

• Campera M, Santini L, Balestri M, Nekaris KAI, Donati G, 'Elevation gradients of lemur abundance emphasise the importance of Madagascar’s lowland rainforest for the conservation of endemic taxa'
  Mammal Review 50 (1) (2020) pp.25-37
  ISSN: 0305-1838
  Abstract
  1. Elevation gradients correlate with changes in several environmental conditions and are known to be related to animal abundance. Animals in regions with a naturally limited extent of lowland rainforest are expected to have evolved adaptations to intermediate elevations that provided a stable environment during their evolution.
  2. Since the lowland rainforest of Madagascar has a limited extent and suffers from increasing anthropogenic pressure, it is essential to understand how well species tolerate intermediate and high elevations. In this study, we aim to quantify the relationship between lemur abundance and elevation in the eastern rainforest of Madagascar.
  3. We correlated abundance data on 26 lemur species (10 genera), including 492 records from 26 studies, with elevation. We analysed the consistency of correlations across species with a meta‐analytical approach. We controlled for species’ body mass, elevational range and median elevation. We then ran generalised linear mixed models to determine whether lemur abundance was related to elevation, body mass, plant productivity and anthropogenic disturbance.
  4. Overall, the abundance of lemur species in Malagasy rainforests was negatively correlated with elevation, and species occupying broader elevational ranges showed stronger correlations. Body mass was not related to species’ tolerance of high elevations. Even though several lemur species are able to occupy the entire elevation gradient, the few remaining patches of lowland rainforests host lemur species at greater abundances than other sites. Abundance across species was negatively related to body mass, elevation and seasonality in plant productivity and positively related to plant productivity.
  5. Despite the ecological flexibility of many lemur species, the remnant patches of lowland rainforests host the highest levels of lemur abundance and are key to lemur conservation. It is crucial to preserve this priority habitat both for biodiversity conservation and for our understanding of lemur adaptations.
  Website 
• Birot H, Campera M, Imron MA, Nekaris KAI, 'Artificial canopy bridges improve connectivity in fragmented landscapes: the case of Javan slow lorises in an agroforest environment'
  American Journal of Primatology (2019)
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract

  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 2016-2017. Motion triggered camera traps collected data for 195 ± SD 85 days on each bridge. We collected 341.76 hours (179.67 h before and 162.09 h 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 instalment mainly for travelling. Slow lorises showed a trend towards 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/h before, 0.43 s/h; 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 travelling 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.
  

  Website 
• Fox, G., R. F. Preziosi, R. E. Antwis, M. Benavides‐Serrato, F. J. Combe, W. Edwin Harris, I. R. Hartley, A. C. Kitchener, S. R. de Kort & A. I. Nekaris, 'Multi-individual Microsatellite identification: a multiple genome approach to microsatellite design (MiMi)'
  Molecular Ecology Resources (2019)
  ISSN: 1755-098X eISSN: 1755-0998
  Abstract

  Bespoke microsatellite marker panels are increasingly affordable and tractable to researchers and conservationists. The rate of microsatellite discovery is very high within a shotgun genomic data set, but extensive laboratory testing of markers is required for confirmation of amplification and polymorphism. By incorporating shotgun next‐generation sequencing data sets from multiple individuals of the same species, we have developed a new method for the optimal design of microsatellite markers. This new tool allows us to increase the rate at which suitable candidate markers are selected by 58% in direct comparisons and facilitate an estimated 16% reduction in costs associated with producing a novel microsatellite panel. Our method enables the visualisation of each microsatellite locus in a multiple sequence alignment allowing several important quality checks to be made. Polymorphic loci can be identified and prioritised. Loci containing fragment‐length‐altering mutations in the flanking regions, which may invalidate assumptions regarding the model of evolution underlying variation at the microsatellite, can be avoided. Priming regions containing point mutations can be detected and avoided, helping to reduce sample‐site‐marker specificity arising from genetic isolation, and the likelihood of null alleles occurring. We demonstrate the utility of this new approach in two species: an echinoderm and a bird. Our method makes a valuable contribution towards minimising genotyping errors and reducing costs associated with developing a novel marker panel. The Python script to perform our method of multi‐individual microsatellite identification (MiMi) is freely available from GitHub (https://github.com/graemefox/mimi).

  Website 
• Geerah RD, O'Hagan RP, Wirdateti W, Nekaris KAI, 'The use of ultrasonic communication to maintain social cohesion in the Javan slow loris (Nycticebus javanicus)'
  Folia Primatologica 90 (2019) pp.392-403
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract

  Only a handful of primate taxa use ultrasonic vocalisations (those ≥20 kHz) to communicate. The extent and uses of ultrasonic communication remain poorly understood, potentially ranging from echolocation, advertisement of reproductive status and resource availability, social cohesion, to predator avoidance. Here, using active acoustics whereby the study subjects were observed throughout their activity period, we describe the first purely ultrasonic call from a strepsirrhine primate (family Lorisidae), recorded in a completely wild setting, and hypothesise about its function. We identified one type of ultrasonic call, the doublet click, from 14 Javan slow lorises (Nycticebus javanicus) produced by males and females of juvenile, subadult and adult ages within their social groups (n = 791, mean = 46.0 kHz). We ran quadratic discriminant function analysis, finding dominant frequency and doublet click duration as the key parameters for identifying individuals’ sex and age. Significantly more vocalisations were produced during affiliative social behaviour, suggesting that the call serves a social cohesion function. Considering the range of other cryptic behaviours, including slow and silent locomotion, and the high degree of territoriality associated with venomous attacks on conspecifics, the call may also serve as a safety strategy, allowing family members to regulate distance from other slow lorises and to communicate cryptically whilst avoiding predators.
  

  Website 
• Reinhardt Kathleen D., Vyazovskiy Vladyslav V., Adriana Hernandez-Aguilar R., Imron Muhammad Ali, Nekaris K. Anne-Isola, 'Environment shapes sleep patterns in a wild nocturnal primate'
  SCIENTIFIC REPORTS 9 (2019)
  ISSN: 2045-2322
  Abstract

  Among primates, the suborder Haplorhini is considered to have evolved a consolidated monophasic sleep pattern, with diurnal species requiring a shorter sleep duration than nocturnal species. Only a few primate species have been systematically studied in their natural habitat where environmental variables, including temperature and light, have a major influence on sleep and activity patterns. Here we report the first sleep study on a nocturnal primate performed in the wild. We fitted seven wild Javan slow lorises (Nycticebus javanicus) in West Java, Indonesia with accelerometers that collected activity data, and installed climate loggers in each individual’s home range to collect ambient temperature readings (over 321 days in total). All individuals showed a strictly nocturnal pattern of activity and displayed a striking synchronisation of onset and cessation of activity in relation to sunset and sunrise. The longest consolidated rest episodes were typically clustered near the beginning and towards the end of the light period, and this pattern was inversely related to daily fluctuations of the ambient temperature. The striking relationship between daily activity patterns, light levels and temperature suggests a major role of the environment in shaping the daily architecture of waking and sleep. We concluded that well-known phenotypic variability in daily sleep amount and architecture across species may represent an adaptation to changes in the environment. Our data suggest that the consolidated monophasic sleep patterns shaped by environmental pressures observed in slow lorises represent phylogenetic inertia in the evolution of sleep patterns in humans.
  

  Website 
• Padfield, Hansen, Davies, Ehrensperger, Slade, Evers, Papargyropoulou, Bessou, Abdullah, Page, Ancrenaz, Aplin, Dzulkafli, Barclay, Chellaiah, Choudhary, Conway, Cook, Copeland, Campos-Arceiz, Deere, Drew, Gilvear, Gray, Haller, Hood, Huat, Huynh, Kangayatkarasu, Koh, Kolandai, Lim, Yeong, Lucey, Luke, Mitchell, Montefrio, Mullin, Nainar, Nekaris, Nijman, Nunes, Nurhidayu, O'Reilly, Puan, Ruppert, Salim, Schouten, Tallontire, Smith, Tao, Tham, Varkkey, Wadey, Yule, Azhar, Sayok, Vairappan, Bicknell and Struebig, 'Co-producing a Research Agenda for Sustainable Palm Oil'
  Science Advances 2 (2019)
  ISSN: 2375-2548
  Abstract

  The rise of palm oil as the world’s most consumed vegetable oil has coincided with exponential growth in palm oil research activity. Bibliometric analysis of research outputs reveals a distinct imbalance in the type of research being undertaken, notably a disproportionate focus on biofuel and engineering topics. Recognizing the expansion of oil palm agriculture across the tropics and the increasing awareness of environmental, social, and economic impacts, we seek to reorientate the existing research agenda toward one that addresses the most fundamental and urgent questions defined by the palm oil stakeholder community. Following consultation with 659 stakeholders from 38 countries, including palm oil growers, government agencies, non-governmental organizations, and researchers, the highest priority research questions were identified within 13 themes. The resulting 279 questions, including 26 ranked as top priority, reveal a diversity of environmental and social research challenges facing the industry, ranging from the ecological and ecosystem impacts of production, to the livelihoods of plantation workers and smallholder communities. Analysis of the knowledge type produced from these questions underscores a clear need for fundamental science programmes, and studies that involve the consultation of non-academic stakeholders to develop “transformative” solutions to the oil palm sector. Stakeholders were most aligned in their choice of priority questions across the themes of policy and certification related themes, and differed the most in environmental feedback, technology and smallholder related themes. Our recommendations include improved regional academic leadership and coordination, greater engagement with private and public stakeholders in Africa, and Central and South America, and enhanced collaborative efforts with researchers in the major consuming countries of India and China.

  Website 
• Campera M, Balestri M, Chimienti M, Nijman V, Nekaris KAI, Donati G, 'Temporal niche separation between the two ecologically similar nocturnal primates Avahi meridionalis and Lepilemur fleuretae'
  Behavioral Ecology and Sociobiology 73 (2019)
  ISSN: 0340-5443 eISSN: 1432-0762
  Abstract

  Time is considered a resource in limited supply, and temporal niche separation is one of the most common strategies that allow ecologically similar species to live in sympatry. Mechanisms of temporal niche separation are understudied especially in cryptic animals due to logistical problems in gathering adequate data. Using high-frequency accelerometers attached to radio-collars, we investigated whether the ecologically similar lemurs Avahi meridionalis and Lepilemur fleuretae in the lowland rainforest of Tsitongambarika, south-eastern Madagascar, show temporal niche separation. Accelerometers stored data with a frequency of 1 Hz for a total of 71 days on three individuals of A. meridionalis and three individuals of L. fleuretae. We extrapolated motor activity patterns via the unsupervised learning algorithm expectation maximisation and validated the results with systematic behavioural observations. Avahi meridionalis showed peaks of activity at twilights with low but consistent activity during the day, while L. fleuretae exhibited more activity in the central hours of the night. Both lemur species had their activity pattern entrained by photoperiodic variations. The pair-living A. meridionalis was found to be lunarphilic while the solitary-living L. fleuretae was lunarphobic. We suggest that these activity differences were advantageous to minimise feeding competition, as an anti-predator strategy, and/or for dietary-related benefits. These findings demonstrate a fine-tuned temporal partitioning in sympatric, ecologically similar lemur species and support the idea that an activity spread over the 24-h, defined here as cathemerality sensu lato, is more common than previously thought in lemurs.
  

  Time is considered a resource in limited supply, and temporal niche separation is one of the most common strategies that allow ecologically similar species to live in sympatry. Mechanisms of temporal niche separation are understudied especially in cryptic animals due to logistical problems in gathering adequate data. Using high-frequency accelerometers attached to radio-collars, we investigated whether the ecologically similar lemurs Avahi meridionalis and Lepilemur fleuretae in the lowland rainforest of Tsitongambarika, south-eastern Madagascar, show temporal niche separation. Accelerometers stored data with a frequency of 1 Hz for a total of 71 days on three individuals of A. meridionalis and three individuals of L. fleuretae. We extrapolated motor activity patterns via the unsupervised learning algorithm expectation maximisation and validated the results with systematic behavioural observations. Avahi meridionalis showed peaks of activity at twilights with low but consistent activity during the day, while L. fleuretae exhibited more activity in the central hours of the night. Both lemur species had their activity pattern entrained by photoperiodic variations. The pair-living A. meridionalis was found to be lunarphilic while the solitary-living L. fleuretae was lunarphobic. We suggest that these activity differences were advantageous to minimise feeding competition, as an anti-predator strategy, and/or for dietary-related benefits. These findings demonstrate a fine-tuned temporal partitioning in sympatric, ecologically similar lemur species and support the idea that an activity spread over the 24-h, defined here as cathemerality sensu lato, is more common than previously thought in lemurs.

  Significance statement
  

  Ecologically similar species may exhibit temporal niche partitioning and separate their peaks of activity when co-occurring in an area. We show for the first time that the mainly nocturnal genus Avahi can exhibit high crepuscular activity with low but consistent bouts of activity (up to 44.6% of daily activity) during the day. We defined this activity as cathemerality sensu lato as opposed to the cathemerality sensu stricto observed in Eulemur sp. We suggest that this flexible activity may be advantageous for the species to minimise feeding competition and predatory pressure, and/or to provide dietary-related benefits. This finding in the secondary nocturnal genus Avahi supports the idea that activity patterns in lemurs are graded and traditional categorisations are inadequate.

  Website 
• Cabana F., Clayton J. B., Nekaris, K. A. I, Wirdateti W., Knights D., Seedorf H., 'Nutrient-based diet modifications impact on the gut microbiome of the Javan slow loris (Nycticebus javanicus)'
  SCIENTIFIC REPORTS 9 (2019)
  ISSN: 2045-2322
  Abstract

  Environment and diet are key factors which shape the microbiome of organisms. There is also a disparity between captive and wild animals of the same species, presumably because of the change in diet. Being able to reverse the microbiome to the wild type is thus particularly important for the reintroduction efforts of Critically Endangered animals. The Javan slow loris (Nycticebus javanicus) is a suitable model, being kept in the thousands within rescue centres throughout Southeast Asia. With next-generation sequencing, we show how a naturalistic diet impacts the gut microbiome of captive slow lorises (Primates: Nycticebus). A comparison of the microbiome of wild animals with captive animals that had been fed a standard captive or improved diet reveals strong microbiome differences between wild and captive animals; however, diet changes failed to alter the microbiome of captive populations significantly. Bifidobacterium was the most abundant genus in wild animals (46.7%) while Bacteroides (11.6%) and Prevotella (18.9%) were the most abundant in captive animals fed the captive and improved diets, respectively. Correlation analyses of nutrients with microbial taxa suggest important implications in using nutrition to suppress potential pathogens, with soluble fibre and water-soluble carbohydrates both being associated with opposing microbiome profiles. The improved diet significantly increased microbe diversity, which exemplifies the importance of high fibre diets; however, wild individuals had lower diversity, which contradicts recent studies. Detection of methanogens appeared to be dependent on diet and whether the animals were living in captivity or in the wild. This study highlights the potential of nutrition in modulating the microbiome of animals prior to release. Unexpectedly, the results were not as significant as has been suggested in recent studies.
  

  Website 
• Nekaris KAI, Weldon A, Imron MA, Maynard KQ, Nijman V, Poindexter SA, Morcatty TQ., 'Venom in Furs: Facial Masks as Aposematic Signals in a Venomous Mammal'
  Toxins 11 (2) (2019)
  ISSN: 2072-6651 eISSN: 2072-6651
  Abstract

  The function of colouration in animals includes concealment, communication and signaling, such as the use of aposematism as a warning signal. Aposematism is unusual in mammals, and exceptions help us to understand its ecology and evolution. The Javan slow loris is a highly territorial venomous mammal that has a distinctive facial mask and monochromatic vision. To help understand if they use aposematism to advertise their venom to conspecifics or predators with different visual systems, we studied a population in Java, Indonesia. Using ImageJ, we selected colours from the facial masks of 58 individuals, converted RBG colours into monochromatic, dichromatic and trichromatic modes, and created a contrast index. During 290 captures, we recorded venom secretion and aggressiveness. Using Non-metric Multidimensional Scaling and generalised additive models for location, scale and shape, we found that young slow lorises differ significantly from adults, being both more contrasting and more aggressive, with aggressive animals showing fewer wounds. We suggest aposematic facial masks serve multiple purposes in slow lorises based on age. Change in colouration through development may play a role in intraspecific competition, and advertise toxicity or aggressiveness to competitors and/or predators in juveniles. Aposematic signals combined with intraspecific competition may provide clues to new venomous taxa among mammals.
  

  Website 
• Luhrs AM, Svensson MS, Nekaris KAI, 'Comparative ecology and behaviour of eastern potto Perodicticus ibeanus and central potto P. edwardsi in Angola, Cameroon, Kenya, Nigeria, Rwanda and Uganda'
  Journal of East African Natural History 107 (1) (2018) pp.17-30
  ISSN: 1026-1613 eISSN: 2224-8862
  Abstract

  Comparative behavioural research reveals both intra- and inter-species diversity among primates. Few long-term behavioural studies have been conducted on African nocturnal primates. Here we describe and compare behavioural and ecological observations on two species of pottos (Perodicticus ibeanus and P. edwardsi) across ten sites. We observed a total of 51 P. edwardsi and 28 P. ibeanus. We recorded all 21 postures within an established lorisid ethogram, as well as 42 of 50 behaviours. Eating, locomotion, freezing, resting and sniffing were the most common behaviours. We recorded behaviours not previously described for perodicticines, including bark chewing and unique vocalisations. Three species of pottos are now recognised, with potentially more species to be revealed within this cryptic and nocturnal genus. Although there are similarities among potto species, we show that unique ecological adaptations and behaviours may further elucidate their diversity.
  

  Website 
• Svensson MS, Nekaris KAI, Bearder SK, Bettridge C, Butynski T, Cheyne SM, Das N, de Jong Y, Luhrs AM, Luncz L, Maddock ST, Perkin A, Pimley E, Poindexter SA, Reinhardt KD, Spaan D, Stark DJ, Starr CR, Nijman V, 'Sleep patterns, daytime predation and the evolution of diurnal sleep site selection in lorisiforms'
  American Journal of Physical Anthropology 166 (3) (2018) pp.563-577
  ISSN: 0002-9483 eISSN: 1096-8644
  Abstract Objectives: Synthesize information on sleep patterns, sleep site use, and daytime predation at sleep sites in lorisiforms of Asia and Africa (ten genera, 36 species), and infer patterns of evolution of sleep site selection. Materials and methods: We conducted fieldwork in twelve African and six Asian countries, collecting data on sleep sites, timing of sleep and predation during daytime. We obtained additional information from literature and through correspondence. Using a phylogenetic approach, we establish ancestral states of sleep site selection in lorisiforms and trace their evolution. Results: The ancestral lorisiform was a fur-clinger and used dense tangles and branches/forks as sleep sites. Use of tree holes and nests as sleep sites emerged ~22 Mya (range 17-26 Mya) in Africa, and use of bamboo emerged ~11 (7-14) Mya in Asia and later in Africa. Nests are commonly used by Galagoides, Paragalago, Galago and Otolemur, tree holes by Galago, Paragalago, Sciurocheirus and Perodicticus, tangles by Nycticebus, Loris, Galagoides, Galago, Euoticus, Otolemur, Perodicticus and Arctocebus, and all but Sciurocheirus and Otolemur additionally sleep on branches/forks. Daytime predation may affect sleep site selection and sleep patterns in some species of Nycticebus, Galago, Galagoides, Otolemur and Perodicticus. Most lorisiforms enter their sleep sites around sunrise and leave around sunset; several are active during twilight or, briefly, during daytime. Conclusion: Variations in sleep behavior, sleep patterns and vulnerability to daytime predation provide a window into the variation that was present in sleep in early primates. Overall, lorisiforms use the daytime for sleeping and no species can be classified as cathemeral or polycyclic.Website 
• Poindexter SA, Reinhardt KD, Nijman V, Nekaris KAI, 'Slow lorises (Nycticebus spp.) display evidence of handedness in the wild and in captivity'
  Laterality: Asymmetries of Body, Brain and Cognition (2018)
  ISSN: 1357-650X eISSN: 1464-0678
  Abstract It has been suggested that strepsirrhines (lemurs, lorises, and galagos) retain the more primitive left hand preference, whilst monkeys and apes more regularly display a right hand preference at the individual-level. We looked to address questions of laterality in the slow loris (Nycticebus spp.) using spontaneous observations of seven wild individuals, unimanual tests in six captive individuals, and photos of 44 individuals in a bilateral posture assessing handedness at the individual- and group-level. During the unimanual reach task, we found at the individual-level, only four slow lorises showed a hand use bias (R: 3, L:1), Handedness index (HI) ranged from -0.57-1.00. In the wild unimanual grasp task we found at the individual-level two individual showed a right-hand bias, the HI ranged from -0.19-0.70. The bilateral venom pose showed a trend toward a right hand dominant grip in those photographed in captivity, but an ambiguous difference in wild individuals. There are many environmental constraints in captivity that wild animals do not face, thus data collected in wild settings are more representative of their natural state. The presence of right-handedness in these species suggest that there is a need to re-evaluate the evolution of handedness in primates.Website 
• Gardiner M, Weldon A, Poindexter SA, Gibson N, Nekaris KAI, 'Survey of practitioners handling slow lorises (Primates: Nycticebus): an assessment of the harmful effects of slow loris bites'
  Journal of Venom Research 9 (2018)
  ISSN: ISSN: 2044-0324
  Abstract Slow lorises (Nycticebus spp.) are one of six venomous mammals, and the only known venomous primate. In the wild envenomation occurs mainly during conspecific competition for mates and territory, but may also be used as an application against parasites or for predator defense. Envenomation in humans is documented, with the most extreme accounts detailing near-fatal anaphylactic shock. From September 2016 – August 2017, we received questionnaire responses from 80 wild animal practitioners working with Nycticebus spp. in zoos, rescue centres and in the wild. We identified 54 practitioners who had experience of being bitten or were otherwise affected by slow loris venom, and an additional 26 incomplete entries. No fatalities were reported. Fifteen respondents noted that medical intervention was required, 12 respondents indicated no reaction to being bitten (9 of these indicated they were wearing gloves). Symptoms for those affected
  included: anaphylactic shock, paraesthesia, haematuria, dyspnoea, extreme pain, infection
  and general malaise. Impact of slow loris bites ranged from instantaneous to long-persisting
  complications, and healing time ranged from 1 day to >8 months. Extremities, including hands and arms, were mostly affected from the bites. Six of nine species of slow loris were reported to bite, with N. pygmaeus being the most common in our sample. We make suggestions regarding the use of these highly threatened yet dangerous primates as unsuitable tourist photo props and zoo animal ambassadors. We discuss the medical complications experienced in relation to protein sensitisation, and bacterial pathogenesis. We recommend future work to ascertain the protein content of slow loris venom to aid in enabling mitigation of risks posed. 
• Hofner AN, Jost Robinson CA, Nekaris KAI, 'Preserving Preuss’s red colobus (Piliocolobus preussi); an analysis of hunting and changing perceptions of primates in Ikenge-Bakoko, Cameroon'
  International Journal of Primatology 39 (5) (2018) pp.895-917
  ISSN: 0164-0291 eISSN: 1573-8604
  Abstract The futures of nonhuman primate species and human communities in shared landscapes rely on our ability to engage with and understand the complex histories and multiscalar aspects of human–animal relationships. We use the Critically Endangered Preuss’s red colobus (Piliocolobus preussi) as a case study to examine the important ways in which histories of multiscalar human–primate interactions play out in the village of Ikenge-Bakoko, Korup National Park, Cameroon. We contextualize ethnographic and catchment data from adult men (N = 32) and women (N = 31) within long-term diurnal primate monitoring datasets to better understand the relationships among hunting practices, local perceptions of diurnal primates, populations of P. preussi, and conservation management. Our data indicate a disconnect between local cultural definitions of “hunter” and Western assumptions as to the makeup and nature of this and other categories. We show that such contradictions can have negative outcomes for conservationists seeking to turn the science of establishing accurate off-take rates of prey species into practical management solutions. Using a single village as a focal point, we highlight the importance of an ethnoprimatological approach to understanding the intricate entanglements among conservation histories, subsistence strategies, and human and nonhuman primate lives. The application of ethnoprimatology is critical for twenty-first century primatologists who must navigate conservation concerns while also acknowledging and valuing the experiences of the human communities living alongside the primates we study.Website 
• Cabana F, Dierenfeld F, Wirdateti, Donati G, Nekaris KAI, 'Trialling nutrient recommendations for slow lorises (Nycticebus spp.) based on wild feeding ecology'
  Journal of Animal Physiology and Animal Nutrition 102 (1) (2018) pp.e1-e10
  ISSN: 0931-2439 eISSN: 1439-0396
  Abstract Slow loris (Nycticebus spp.) captive diets have been based on routine and anecdotes rather than scientific fact. The growing body of evidence contradicts the high fruit diet supported by such anecdotes. Non-human primate nutrient requirements are grouped into new (based on the common marmoset Callithrix jacchus) or old world (based on rhesus macaques Macaca mulatta) primates. Slow lorises are known to suffer from many health ailments in captivity such as dental disease, obesity, wasting and kidney issues all of which have been linked to diet. This study aimed to estimate nutrient intake from free-ranging slow lorises and to determine whether this intake can be used as nutrient recommendations. We collected data of nutrient intake, food passage rate and digestibility of captive slow lorises on three diet treatments 1: current captive type diet which is mostly fruits, 2: wild-type diet made only of food items from their natural diet, 3: new diet made to reflect wild slow loris nutrient intake. In order to validate our nutrient recommendations, diets 2 and 3 would have to be significantly different to Diet 1 in terms of nutrients, but not different from each other. Captive diets were significantly higher in soluble carbohydrates and lower in minerals and fibre fractions than both diets 2 and 3. Diets 2 and 3 led to a significantly increased food passage time and to more effective fibre and calcium digestion compared to Diet 1. We also observed obese individuals lost weight and underweight individuals gained weight. Our nutrient recommendations have been validated by our trials, and new or old world monkey nutrient recommendations are not consistent with our results. Diets should be high in protein and fibre and low in soluble carbohydrates and fats.Website 
• Fuller G, Eggen WF, Wirdateti W, Nekaris KAI, 'Welfare impacts of the illegal wildlife trade in a cohort of confiscated greater slow lorises, Nycticebus coucang'
  Journal of Applied Animal Welfare Science 21 (3) (2017) pp.224-238
  ISSN: 1088-8705 eISSN: 1532-7604
  Abstract 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.Website 
• Donati G, Santini L, Eppley TM, Arrigo-Nelson SJ, Balestri M, Boinski S, Bollen A, Bridgeman LL, Campera M, Carrai V, Chalise MK, Derby Lewis A, Hohmann G, Kinnaird MF, Koenig A, Kowalewski M, Lahann P, McLennan MR, Nekaris AKI, Nijman V, Norscia I, Ostner J, Polowinsky SY, Schülke O, Schwitzer C, Stevenson PR, Talebi MG, Tan C, Tomaschewski I, Vogel ER, Wright PC, Ganzhorn JU, 'Low levels of fruit nitrogen as drivers for the evolution of Madagascar’s primate communities'
  Scientific Reports 7 (2017)
  ISSN: 2045-2322
  Abstract The uneven representation of frugivorous mammals and birds across tropical regions – high in the New World, low in Madagascar and intermediate in Africa and Asia – represents a long-standing enigma in ecology. Several hypotheses have been proposed to explain these differences but the ultimate drivers remain unclear. Here, we tested the idea that fruits in Madagascar contain insufficient nitrogen to meet primate metabolic requirements, thus constraining the evolution of frugivory. We performed a global analysis of nitrogen in fruits consumed by primates, as collated from 79 studies. Our results showed that average frugivory among lemur communities was lower compared to New World and Asian-African primate communities. Fruits in Madagascar contain lower average nitrogen than those in the New World and Old World. Nitrogen content in the overall diets of primate species did not differ significantly between major taxonomic radiations. There is no relationship between fruit protein and the degree of frugivory among primates either globally or within regions, with the exception of Madagascar. This suggests that low protein availability in fruits influences current lemur communities to select for protein from other sources, whereas in the New World and Old World other factors are more significant in shaping primate communities.Website 
• Cabana F, Dierenfeld E, Wirdateti W, Donati G, Nekaris KAI, 'Slow lorises (Nycticebus spp.) really are slow: A study into food passage rates'
  International Journal of Primatology 38 (5) (2017) pp.900-913
  ISSN: 0164-0291 eISSN: 1573-8604
  Abstract 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 fibre 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 four N. menegensis) in an Indonesian rescue centre and four captive-born slow lorises (two N. bengalensis and two N. pygmaeus) in a UK institution. We fed the Indonesian animals 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 animals a diet of gum, vegetables, insects and hard-boiled eggs. We formulated this diet to mimic the wild diet, with notably higher fibre 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 prior to their main diet. We noted the date and time of feeding and of appearances of the marker in faeces. We weighed food given and left over for each animal 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. pygmaeus may have the slowest MRT of all primates in relation to body mass. The digestive flexibility of exudativorous primates should allow them to maximise fermentation opportunities when they ingest more (appropriate) fibre by increasing the amount of time the fibre 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.Website 
• Nekaris KAI, McCabe S, Spaan D, Ali MI, Nijman V, 'A novel application of cultural consensus models to evaluate conservation education programs'
  Conservation Biology 31 (2017)
  ISSN: 0888-8892 eISSN: 1523-1739
  Abstract Conservation professionals recognize the need to evaluate education initiatives with a flexible
  approach that is culturally appropriate. Cultural-consensus theory (CCT) provides a
  framework for measuring the extent to which beliefs are communally held and has long been
  applied by social scientists. In a conservation-education context, we applied CCT and used
  free lists (i.e., a list of items on a topic stated in order of cultural importance) and domain
  analysis (analysis of how free lists go together within a cultural group) to evaluate a
  conservation education program in which we used a children’s picture book to increase
  knowledge about and empathy for a critically endangered mammal, the Javan slow loris
  (Nycticebus javanicus). We extracted free lists of keywords generated by students (n=580 in
  18 schools) from essays they wrote before and after the education program. In 2 classroom
  sessions conducted approximately 18 weeks apart, we asked students to write an essay about
  their knowledge of the target species and then presented a book and several activities about
  slow loris ecology. Prior to the second session, we asked students to write a second essay. We
  generated free lists from both essays, quantified salience of terms used, and conducted
  minimal residuals factor analysis to determine presence of cultural domains surrounding slow
  lorises in each session. Students increased their use of words accurately associated with slow
  loris ecology and conservation from 43% in initial essays to 76% in final essays . Domain
  coherence increased from 22% to 47% across schools. Fifteen factors contributed to the
  domain slow loris. Between the first and second essays, factors that showed the greatest
  change were feeding ecology and slow loris as a forest protector, which increased 7-fold, and
  the humancentric factor, which decreased 5-fold. As demonstrated by knowledge retention
  and creation of unique stories and conservation opinions, children achieved all six levels of
  Bloom’s taxonomy of learning domains. Free from the constraints of questionnaires and
  This article is protected by copyright. All rights reserved.
  3
  surveys, CCT methods provide a promising avenue to evaluate conservation education
  programsWebsite 
• Balestri M, Campera M, Nekaris KAI, Donati G, 'Assessment of long-term retention of environmental education lessons given to teachers in rural areas of Madagascar'
  Applied Environmental Education and Communication 16 (4) (2017) pp.298-311
  ISSN: 1533-015X eISSN: 1533-0389
  Abstract Assessing the retention of knowledge is the first step of environmental education programs. The low education level in rural areas is one factor influencing habitat loss in Madagascar. We tested whether environmental education lessons given to teachers from a municipality, Iaboakoho, in a priority area for lemur conservation were retained after one year. Questionnaires were given to teachers from Iaboakoho and from other three neighbouring municipalities (control groups). Teachers from Iaboakoho had higher scores than the others. Knowledge gained was retained and might be transferred to children. Increasing proenvironmental attitudes and behaviours is the next step to reduce environmental exploitation.Website 
• Kitson H, Nekaris KAI, 'Instagram-fuelled illegal slow loris trade uncovered in Marmaris, Turkey'
  Oryx 51 (3) (2017) pp.394-394
  ISSN: 0030-6053 eISSN: 1365-3008
  Website 
• Poindexter SA, Nekaris KAI, 'Vertical clingers and gougers: Rapid acquisition of adult limb proportions facilitates feeding behaviours in young Javan slow lorises (Nycticebus javanicus)'
  Mammalian Biology 87 (Nov. 2017) (2017) pp.40-49
  ISSN: 1616-5047
  Abstract Animals of all ages need to access essential food resources, either on their own or with the assistance of conspecifics. Rapid physical and behavioural development is one strategy to help young animals reach adulthood. Specialized gum-feeding mammals exploit a food type that is relatively difficult to access and digest and must possess the appropriate adaptions to access large vertical substrates, i.e. tree trunks. Unlike other gum feeding mammals, the Javan slow loris (Nycticebus javanicus) lacks physical structures, such as keeled nails or claws, which animals commonly use to secure themselves to large vertical substrates. To understand how slow lorises of all ages exploit gum, we examined their vertical gouging posture, locomotor behaviour, habitat use during feeding, and their morphometric measures across three age classes (adult, sub-adult, juvenile). Using data collected in Cipaganti, Java, Indonesia between April 2012 and April 2016, we found that individuals of N. javanicus rely on their hand, foot, and limb morphology to maintain vertical gouging postures, in place of claws or keeled nails. Locomotor behaviour, position in tree, and tree DBH showed no significant difference across age classes while feeding. Juveniles were indistinguishable from adults and sub-adults in regards to limb proportion indices, lower leg length, hand span and foot span. Some morphometric measures scaled isometrically e.g. arm length, but those highlighted during prolonged vertical postures scaled allometrically e.g. leg, hand, and foot measures. These findings suggest that the rapid behavioural and physical development of key features may act as an ontogenetic adaptation to facilitate access to a stable food resource at a young age. The Javan slow loris exemplifies the complex relationship that exists between an animal’s diet and the specializations that facilitate access to these food resources.Website 
• Nijman V, Nekaris KAI, 'The Harry Potter effect: The rise in trade of owls as pets in Java and Bali, Indonesia'
  Global Ecology and Conservation 11 (2017) pp.84-94
  ISSN: 2351-9894
  Abstract Hundreds of species of wild-caught birds are offered for sale in the bird markets of Java and Bali, Indonesia, to meet the demand for the largely-domestic pet and songbird trade. In the past, owls were offered only in very small numbers in these bird markets but since the release of the Harry Potter series in Indonesia in the early 2000s their popularity as pets has increased. Whereas in the past owls were collective known as Burung Hantu (“Ghost birds”), in the bird markets they are now commonly referred to as Burung Harry Potter (“Harry Potter birds”). We made a retrospective quantitative assessment of the abundance of owls in the bird markets (1979–2010) and conducted 109 surveys in 20 bird markets in 2012–2016 to quantify owls in trade. In the 1980s, 1990s and early 2000s owls were rarely recorded in Indonesia's bird markets, typically one or two and up to five per survey, and frequently no owls were recorded at all. The trade was largely confined to small scops owls. In the late 2000s more species were offered for sale, including barn and bay owls, and larger owl species such as wood-owls, eagle-owls and fish-owls; typically 10 + owls were observed per survey. In recent years, the number of owl species increased even more, and on average we recorded 17 owls per survey, yielding a total of 1810 owls, and in >90% of the surveys owls were present. In the larger bird markets in Jakarta and Bandung typically 30 to 60 owls are on offer of up to 8 species at a time. The number of owls as a proportion of all birds in the markets increased from <0.06% prior to 2002 to >0.43% post 2008, suggesting a delayed Harry Potter effect. Over this period, common species have become cheaper and less common ones have become more expensive. The owls are largely, if not exclusively, wild-caught and are sold into the domestic pet market. The release of Harry Potter films and novels in Indonesia coincided with the rise of the Internet and social media and, with some delay, the emergence of pet owl interest groups on Java and Bali, thus preventing us to demonstrate a causal Harry Potter effect on the owl trade. The overall popularity of owls as pets in Indonesia has risen to such an extent that it may imperil the conservation of some of the less abundant species. Inclusion of owls on Indonesia's protected species list, alongside all diurnal raptors, may be a first step to mitigate the negative effects of this emerging trend.Website 
• Kappeler PM, Cuozzo FP, Fichtel C, Ganzhorn JU, Gursky-Doyen S, Irwin MT, Ichino S, Lawler R, Nekaris KAI, Ramnamanjato JB, Radespiel U, Sauther ML, Wright PC, Zimmermann E, 'Long-term field studies of lemurs, lorises, and tarsiers'
  Journal of Mammalogy 98 (3) (2017) pp.661-669
  ISSN: 1545-1542
  Abstract Lemurs, lorises, and tarsiers are socially and ecologically diverse primates that include some of the most endangered mammals. We review results of long-term studies of 15 lemur species from 7 sites in Madagascar and 1 species each of loris and tarsier in Indonesia. We emphasize that the existence of long-term study populations is a crucial prerequisite for planning and conducting shorter studies on specific topics, as exemplified by various ecophysiological studies of lemurs. Extended studies of known individuals have revealed variation in social organization within and between ecologically similar species. Even for these primates with relatively fast life histories, it required more than a decade of paternity data to characterize male reproductive skew. The long-term consequences of female rank on reproductive success remain poorly known, however. Long-term monitoring of known individuals is the only method to obtain data on life-history adaptations, which appear to be shaped by predation in the species covered here; long-term studies are also needed for addressing particular questions in community ecology. The mere presence of long-term projects has a positive effect on the protection of study sites, and they generate unique data that are fundamental to conservation measures, such as close monitoring of populations.
  
  Los lémures, lorises y tarseros son grupos de primates, muy diversos social y ecológicamente, que incluyen algunas de las especies de mamíferos más amenazadas. Se revisaron los resultados de estudios a largo plazo de 15 especies de lémures en 7 áreas de estudio en Madagascar y una especie de loris y otra de tarsero, en Indonesia. Se resalta la importancia de las áreas de estudio a largo plazo como prerrequisito esencial para planear y llevar a cabo estudios de menor duración sobre temas especificos, como lo ejemplifican varios estudios eco-fisiológicos en lemurs. Estudios a largo plazo de individuos conocidos, han revelado variacion en la organizacion social dentro y entre especies ecologicamente similares. Incluso en estos primates, con un ciclo de vida relativamente corto, se necesita recopilar más de una década de datos de paternidad para determinar el sesgo reproductivo de los machos. Sin embargo, aún se sabe poco de las consecuencias a largo plazo que la posición de las hembras en la jerarquía social puede tener en su éxito reproductivo. El monitoreo a largo plazo de individuos conocidos es el unico metodo para obtener datos sobre adaptaciones en su history natural, las cuales aparentemente son moduladas en function de la predacion a las especies estudiadas en este trabajo. Estos estudios a largo plazo son tambien necesarios para afrontar temas específicos sobre la ecología de estas comunidades. La simple presencia de proyectos a largo plazo tienen un efecto positive en la proteccion de los lugares de studio, como asi tambien generan datos unicos que son fundamentals para apoyar medidas de conservacion tales como el monitoreo de poblaciones.
  Website 
• Nekaris KAI, Poindexter S, Reinhardt KD, Sigaud M, Cabana F, Wirdateti W, Nijman V, 'Coexistence between Javan Slow Lorises (Nycticebus javanicus) and Humans in a Dynamic Agroforestry Landscape in West Java, Indonesia'
  International Journal of Primatology 38 (2) (2017) pp.303-320
  ISSN: 0164-0291
  Abstract 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.Website 
• Cabana F, Dierenfeld ES, Wirdateti, Donati G, Nekaris KAI, 'Exploiting a readily available but hard to digest resource: A review of exudativorous mammals identified thus far and how they cope in captivity'
  Integrative Zoology 13 (1) (2017) pp.94-111
  ISSN: 1749-4869
  Abstract Gum is a widely available carbohydrate, composed mainly of non-digestible structural carbohydrates. No mammalian enzymes can digest gum, therefore a mammal ingesting gum must rely on microbial fermentation in order to access the energy it possesses. Gums are known as relatively nutrient poor. Despite this, some mammals have evolved to exploit this food resource. We aim to review the literature for all mammal species which have been recorded to ingest gum, whether quantified or not and discuss this in context of their evolutionary adaptations. We also investigated the recommended captive diets for these species to look at if gum is recommended. We conducted a literature search on ISI Web of Knowledge to tabulate all mammal species observed ingesting gum and classify them as obligate, facultative or opportunistic feeders. We encountered 94 mammal species which eat gum in the wild (26 obligate feeders, 35 facultative feeders and 33 opportunistic feeders). Obligate feeders have entirely evolved to exploit this resource but were found to not be given gum in captivity, which may explain why they are failing to thrive, as opposed to facultative feeders which have fewer issues. Gum may be necessary for the health of obligate feeders in captivity. Future research should focus on the physiological effects that gum ingestion poses on different digestive systems.Website 
• Svensson MS, Bersacola E, Mills MSL, Munds RA, Nijman V, Perkin A, Masters JC, Couette S, Nekaris KAI, Bearder SK, 'A giant among dwarfs: a new species of galago (Primates: Galagidae) from Angola'
  American Journal of Physical Anthropology 163 (1) (2017) pp.30-43
  ISSN: 0002-9483 eISSN: 1096-8644
  Abstract Objectives. Based on vocalization recordings of an unknown galago species, our main objectives were to compare morphology and call structure with known closely-related taxa and describe a new species of galago.
  Materials and methods. We conducted field surveys in three forest habitats along the escarpment region in western Angola (Kumbira Forest, Bimbe Area, and Northern Scarp Forest), and examined galago specimens from museums worldwide. We digitized and analyzed calls using Avisoft SASLab Pro software. We also compared museum specimens from Angola with other Galago and Galagoides specimens, and conducted comparative analyses (ANOVA and between group principle component analysis) based on a set of twelve linear measurements of skulls and teeth.
  Results. We describe the new species to which we give the name Angolan dwarf galago, Galagoides kumbirensis sp. nov. The new species has a loud and characteristic crescendo call, used by other Galagoides spp. (sensu stricto) in West Africa to attract companions and repel rivals. However, this call shows species-typical differences from its closest relatives. Galagoides kumbirensis sp. nov. is also distinguished by differences in the skull morphology, pelage color and facial markings, as well as a larger body size, similar to that of Galago moholi, which is not known to be sympatric.
  Conclusion. This discovery points to the importance of Angolan forests as refuges for endemic biodiversity. These forests are under severe threat from overexploitation, and there is an urgent need to establish conservation measures and designate protected areas.Website 
• Fuller G, Nijman V, Wirdateti, Nekaris KAI, 'Do chemical cues in the venom of slow lorises repel avian predators?'
  Emu 116 (4) (2017) pp.435-439
  ISSN: 0158-4197 eISSN: 1448-5540
  Abstract Raptors are confirmed predators of Asian slow lorises (Nycticebus spp.), the only primates with a toxic bite. A possible function of this venom is to protect against predators. Slow lorises release volatile chemicals when threatened, thus potentially communicating their venomous status towards predators. Crested Serpent-eagles (Spilornis cheela) and Changeable Hawk-eagles (Nisaetus cirrhatus) are known to prey on venomous snakes and small mammals and are potential predators of slow lorises. We tested the anti-predator potential of slow loris venom by presenting pieces of chicken combined with swabs of Greater Slow Loris (Nycticebus coucang) venom to 10 Changeable Hawk-eagles and 5 Crested Serpent-eagles. The eagles showed few behavioural responses in reaction to slow loris venom, examining swabs with venom or control scents equally. Both eagle species did show higher rates of face-rubbing behaviour following consumption of foods paired with venom compared with control scents. Our data suggest that slow loris venom does not function to repel avian predators, but may have an anti-predator defence function. While the eagles are not repelled by the smell of slow lorises, contact with their venom causes discomfort, potentially limiting the palatability of slow lorises to eagles.Website 
• Cabana F, Dierenfeld E, Wirdateti W, Donati G, Nekaris KAI, 'The seasonal feeding ecology of the Javan slow loris (Nycticebus javanicus)'
  American Journal of Physical Anthropology (2017)
  ISSN: 0002-9483
  Abstract Objectives. To describe the strategy employed by exudativorous primates during seasonal shifts in food abundance using the Javan slow loris as a model. Males and females may cope differently as well as exploit fallback foods in different proportions.
  Materials and methods. Observing 15 free ranging Javan slow lorises over a year, we quantified their seasonal diet and nutrient intake using intake rates. For gum intake rates, we conducted a trial with 10 captive Javan slow lorises measuring the length of time it took for them to ingest 10 g of gum. We monitored phenology in our field site over five plots that were assessed monthly. We weighed our free-ranging animals every six months. We analyzed all food items slow lorises ingested for macronutrients using the nutritional geometry framework.
  Results. The slow loris diet consisted of eight food categories, with gum and insects being the major food sources in terms of wet weight intake. The captive gum trials resulted in an intake rate of 0.021 g/s. All food items eaten by wild Javan slow lorises were available in the wet season and were restricted in the dry season. Males and females reacted differently to seasonal abundances with females ingesting more protein, gum, fruits and flowers and males ingesting more fiber.
  Conclusions. The strategy used by the Javan slow lorises during periods of lower food availability were similar to folivorous primates and included increased dependence on lower quality foods. The reproductive costs of gestation and lactation may place a burden on females that requires them to alter their foraging strategy during the dry season to ensure enough protein and overall energy is ingested. The overall strategy used by these exudativorous primates is one of nutrient maximization as no nutrient was clearly preferred over another.Website 
• Estrada A, Garber P A, Rylands A B, Roos C, Fernandez-Duque E, Di Fiore A , Nekaris K A-I, Nijman V, Heymann E W , Lambert J E, Rovero F, Barelli C, Setchell J M, Gillespie T R, Mittermeier R A, Verde Arregoitia L, de Guinea M, Gouveia S, Dobrovolski R, Shanee S, Shanee N, Boyle S A, Fuentes A, MacKinnon K C, Amato K R, Meyer A L S, Wich S, Sussman R W, Pan R, Kone I, Li B, 'Impending extinction crisis of the world’s primates: Why primates matter'
  Science Advances 3 (1) (2017)
  ISSN: 2375-2548
  Abstract Non-human primates, our closest biological relatives, play important roles in the livelihoods, cultures and religions of many societies, and offer unique insights into human evolution, biology, behavior and the threat of emerging diseases. They are an essential component of tropical biodiversity, contributing to forest regeneration and ecosystem health. The most recent compilation of primate taxonomy lists 504 species, 197 subspecies and 79 genera distributed in the Neotropics, mainland Africa, Madagascar and Asia. Alarmingly, ~60% of primate species are now threatened with extinction as a result of unsustainable human activities, including illegal hunting and those resulting in extensive land-cover changes: industry driven agricultural production, deforestation, livestock and cattle ranching, oil and gas drilling, mining, dam building, climate change, and poor governance. Although drivers of primate decline vary by region, it is clear that decreasing the per capita demand of industrialized nations, lowering human birth rates and population growth, improving health, reducing poverty and gender biases in education, developing sustainable land-use initiatives, and preserving traditional livelihoods in primate range countries are all part of a comprehensive solution. Despite the existing threats to primate survival, we are adamant that primate conservation is not yet a lost cause. We still have the opportunity to reduce the human impact to primates and their habitats, but that demands raising greater local, regional and global public awareness of the plight of the world’s primates and the costs of their loss to ecosystem health, human culture and ultimately human survival.
• Svensson M S, Shanee S, Shanee N, Bannister FB, Cervera L, Donati G, Huck M, Jerusalinsky L, Juarez CP, Maldonado AM, Martinez Molinedo J, Méndez-Carvajal PG, Molina Argandoña MA, Mollo Vino A D, Nekaris KAI, Peck M, Rey-Goyeneche J, Spaan D, Nijman V, 'Disappearing in the night: an overview on trade and legislation of night monkeys in South and Central America'
  Folia Primatologica 87 (5) (2017) pp.332-348
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract The international trade in night monkeys (Aotus spp.), found throughout Central and South America, has been regulated by the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) since 1975. We present a quantitative analysis of this trade from all 9 range countries, over 4 decades, and compare domestic legislation to CITES regulations. Night monkeys were exported from 8 of the 9 habitat countries, totalling 5,968 live individuals and 7,098 specimens, with trade of live individuals declining over time. In terms of species, the most commonly traded was Aotus nancymaae (present in Brazil, Colombia, Peru) followed by A. vociferans (Brazil, Colombia, Ecuador, Peru) and A. zonalis (Colombia, Panama). There was no significant correlation between levels of trade and species' geographic range size or the number of countries in which a species occurs. Five countries have legislation that meets CITES requirements for implementation, whereas the other 4 countries' legislation showed deficiencies. Research conducted in Colombia, Peru, and Brazil suggests significant cross-border trade not captured in official international trade registers. Although international trade has diminished, current trends suggest that populations of rarer species may be under unsustainable pressure. Further research is needed to quantify real trade numbers occurring between habitat countries.Website 
• Reinhardt KD, Wirdateti, Nekaris KAI, 'Climate-mediated activity of the Javan slow loris, Nycticebus javanicus. AIMS Environmental Science'
  AIMS Environmental Science 3 (2) (2016) pp.249-260
  ISSN: 2372-0344 eISSN: 2372-0352
  Abstract Joint impacts of anthropogenic disturbance and climate change are of pressing concern for modern conservationists. Climate change patterns have various diminishing effects on the biodiversity of an ecosystem, requiring an understanding of a species’ ability to adapt. Agricultural practices are expanding at an altitudinal gradient on the Indonesian island of Java, forcing endemic species to range at increased elevation with lower temperatures, and in human-populated areas. One example is the Critically Endangered Javan slow loris (Nycticebus javanicus), which finds itself increasingly restricted to montane regions with extreme climate patterns and habitat disturbance. We observed wild N. javanicus in a highly fragmented, montane agroforest area to determine if climate variables and forest connectivity influence activity budget and behavior. Lorises ranged at different altitudes (1275 m above sea level (asl)—1570 m asl) and were observed for six months in Cipaganti, West Java. Using multinomial regression analyses, we found loris individuals were most likely to engage in increased foraging, feeding and travelling behavior than resting when relative humidity increases and in habitats with greater forest connectivity. Regression analyses found effects of relative humidity and forest connectivity to be the most significant predictors of N. javanicus foraging behavior (P = 0.001, P = 0.030). We suggest that future-climate shifts and increased anthropogenic disturbance will detrimentally influence wild populations of N. javanicus, requiring immediate plans for mitigation in conserving these already scarce wild populations. We also suggest the altering of reintroduction protocols in relation to climate and geographic region.Website 
• Nekaris KAI, Musing L, Vazquez AG, Donati G, 'Is tickling torture? Assessing welfare towards slow lorises (Nycticebus spp) within Web 2.0 videos'
  Folia Primatologica 86 (6) (2016) pp.534-551
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract Videos, memes and images of pet slow lorises have become increasingly popular on the Internet. Although some video sites allow viewers to tag material as ‘animal cruelty', no site has yet acknowledged the presence of cruelty in slow loris videos. We examined 100 online videos to assess whether they violated the ‘five freedoms' of animal welfare and whether presence or absence of these conditions contributed to the number of thumbs up and views received by the videos. We found that all 100 videos showed at least 1 condition known as negative for lorises, indicating absence of the necessary freedom; 4% showed only 1 condition, but in nearly one third (31.3%) all 5 chosen criteria were present, including human contact (57%), daylight (87%), signs of stress/ill health (53%), unnatural environment (91%) and isolation from conspecifics (77%). The public were more likely to like videos where a slow loris was kept in the light or displayed signs of stress. Recent work on primates has shown that imagery of primates in a human context can cause viewers to perceive them as less threatened. Prevalence of a positive public opinion of such videos is a real threat towards awareness of the conservation crisis faced by slow lorises.Website 
• Nijman V, Spaan D, Nekaris KAI, 'Large-scale trade in legally protected marine mollusc shells from Java and Bali, Indonesia'
  PLoS ONE 10 (12) (2015)
  ISSN: 1932-6203
  Abstract Background
   
  Tropical marine molluscs are traded globally. Larger species with slow life histories are under threat from over-exploitation. We report on the trade in protected marine mollusc shells in and from Java and Bali, Indonesia. Since 1987 twelve species of marine molluscs are protected under Indonesian law to shield them from overexploitation. Despite this protection they are traded openly in large volumes.
   
  Methodology/Principal Findings
   
  We collected data on species composition, origins, volumes and prices at two large open markets (2013), collected data from wholesale traders (2013), and compiled seizure data by the Indonesian authorities (2008–2013). All twelve protected species were observed in trade. Smaller species were traded for <USD1.00 whereas prices of larger species were USD15.00–40.00 with clear price-size relationships. Some shells were collected locally in Java and Bali, but the trade involves networks stretching hundreds of kilometres throughout Indonesia. Wholesale traders offer protected marine mollusc shells for the export market by the container or by the metric ton. Data from 20 confiscated shipments show an on-going trade in these molluscs. Over 42,000 shells were seized over a 5-year period, with a retail value of USD700,000 within Indonesia; horned helmet (Cassis cornuta) (>32,000 shells valued at USD500,000), chambered nautilus (Nautilus pompilius) (>3,000 shells, USD60,000) and giant clams (Tridacna spp.) (>2,000 shells, USD45,000) were traded in largest volumes. Two-thirds of this trade was destined for international markets, including in the USA and Asia-Pacific region.
   
  Conclusions/Significance
   
  We demonstrated that the trade in protected marine mollusc shells in Indonesia is not controlled nor monitored, that it involves large volumes, and that networks of shell collectors, traders, middlemen and exporters span the globe. This impedes protection of these species on the ground and calls into question the effectiveness of protected species management in Indonesia; solutions are unlikely to be found only in Indonesia and must involve the cooperation of importing countries.
  Website 
• Nijman V, Spaan D, Rode-Margono EJ, Wirdateti, Nekaris KAI, 'Changes in the primate trade in Indonesian wildlife markets over a 25-year period: fewer apes and langurs, more macaques and slow lorises'
  American Journal of Primatology 79 (11) (2015)
  ISSN: 0275-2565
  Abstract Indonesia has amongst the highest primate species richness, and many species are included on the country's protected species list, partially to prevent over-exploitation. Nevertheless traders continue to sell primates in open wildlife markets especially on the islands of Java and Bali. We surveyed 13 wildlife markets in 2012–2014 and combined our results with previous surveys from 1990–2009 into a 122-survey dataset with 2,424 records of 17 species. These data showed that the diversity of species in trade decreased over time, shifting from rare rainforest-dwelling primates traded alongside more widespread species that are not confined to forest to the latter type only. In the 1990s and early 2000s orangutans, gibbons and langurs were commonly traded alongside macaques and slow lorises but in the last decade macaques and slow lorises comprised the bulk of the trade. In 2012–2014 we monitored six wildlife markets in Jakarta, Bandung and Garut (all on Java), and Denpasar (Bali). During 51 surveys we recorded 1,272 primates of eight species. Traders offered long-tailed macaque (total 1,007 individuals) and three species of slow loris (228 individuals) in five of the six markets, whereas they traded ebony langurs (18 individuals), and pig-tailed macaques (14 individuals) mostly in Jakarta. Pramuka and Jatinegara markets, both in Jakarta, stood out as important hubs for the primate trade, with a clear shift in importance over time from the former to the latter. Slow lorises, orangutans, gibbons and some langurs are protected under Indonesian law, which prohibits all trade in them; of these protected species, only the slow lorises remained common in trade throughout the 25-year period. Trade in non-protected macaques and langurs is subject to strict regulations—which market traders did not follow—making all the market trade in primates that we observed illegal. Trade poses a substantial threat to Indonesian primates, and without enforcement, the sheer volume of trade may mean that species of Least Concern or Near Threatened may rapidly decline.Website 
• Svensson MS, Ingram DJ, Nekaris KAI, Nijman V, 'Trade and ethnozoological use of African lorisiforms in the last 20 years'
  Hystrix the Italian Journal of Mammalogy 26 (2) (2015) pp.153-161
  ISSN: 0394-1914 eISSN: 1825-5272
  Abstract Trade in primates is considered a major impediment to primate conservation globally. The bushmeat trade in West and Central Africa is considered largely unsustainable and represents one of the main threats to biodiversity. Furthermore, the use of primates in traditional practices and medicine includes a third of the African primate species. Little is known about the trade in the African mainland lorisiforms; pottos, angwantibos and galagos. Aiming to fill this knowledge gap we created an online survey, conducted a literature review, and analyzed CITES trade records, focusing on the last two decades. We obtained 188 questionnaire responses from researchers and people working in 31 different countries in sub-Saharan Africa. We found a total of 33 publications reporting on trade in African lorisiforms, and CITES records indicate that almost 2000 lorisiforms were traded internationally from African range countries. Fifty-three percent of respondents provided meaningful details about aspects of the trade in African lorisiforms from 50% of the range countries. Galagos were reported by respondents in larger numbers than pottos and angwantibos, and mainly occurred in the pet trade. Pottos were the most frequently mentioned taxon in the literature, when all trade types were combined. Across all of the sources (online survey, literature and CITES database), trade in pottos and angwantibos was reported from 12 countries, and galagos from 23 countries. Trade was reported to occur mainly within rural settings (64%), potentially indicating that demand is not high enough to fuel long distance trading. However, as seen in the Asian lorisiforms, once quantitative studies were conducted, the threat that trade posed became alarmingly apparent and is now considered a major impediment to their conservation. Our insight into the trade of African lorisiforms should be followed up with concerted studies, with an emphasis on quantifying trade to  the species level.Website 
• Rode-Margono JE, Nekaris KAI, 'Cabinet of curiosities: Venom systems and their ecological function in mammals, with a focus on primates'
  Toxins 7 (7) (2015) pp.2639-2658
  ISSN: 2072-6651 eISSN: 2072-6651
  Abstract Venom delivery systems (VDS) are common in the animal kingdom, but rare amongst mammals. New definitions of venom allow us to reconsider its diversity amongst mammals by reviewing the VDS of Chiroptera, Eulipotyphla, Monotremata, and Primates. All orders use modified anterior dentition as the venom delivery apparatus, except Monotremata, which possesses a crural system. The venom gland in most taxa is a modified submaxillary salivary gland. In Primates, the saliva is activated when combined with brachial gland exudate. In Monotremata, the crural spur contains the venom duct. Venom functions include feeding, intraspecific competition, anti-predator defense and parasite defense. Including mammals in discussion of venom evolution could prove vital in our understanding protein functioning in mammals and provide a new avenue for biomedical and therapeutic applications and drug discovery.Website 
• Pozzi L, Nekaris KAI, Perkin A, Bearder SK, Pimley ER, Schulze H, Streicher U, Zischler H, Bruford M, Zinner D, Roos C, 'Remarkable Ancient Divergences Amongst Neglected Lorisiform Primates'
  Zoological Journal of the Linnean Society 175 (3) (2015) pp.661-674
  ISSN: 0024-4082 eISSN: 1096-3642
  Abstract Lorisiform primates (Primates: Strepsirrhini: Lorisiformes) represent almost 10% of the living primate species and are widely distributed in sub-Saharan Africa and South/South-East Asia; however, their taxonomy, evolutionary history, and biogeography are still poorly understood. In this study we report the largest molecular phylogeny in terms of the number of represented taxa. We sequenced the complete mitochondrial cytochrome b gene for 86 lorisiform specimens, including ∼80% of all the species currently recognized. Our results support the monophyly of the Galagidae, but a common ancestry of the Lorisinae and Perodicticinae (family Lorisidae) was not recovered. These three lineages have early origins, with the Galagidae and the Lorisinae diverging in the Oligocene at about 30 Mya and the Perodicticinae emerging in the early Miocene. Our mitochondrial phylogeny agrees with recent studies based on nuclear data, and supports Euoticus as the oldest galagid lineage and the polyphyletic status of Galagoides. Moreover, we have elucidated phylogenetic relationships for several species never included before in a molecular phylogeny. The results obtained in this study suggest that lorisiform diversity remains substantially underestimated and that previously unnoticed cryptic diversity might be present within many lineages, thus urgently requiring a comprehensive taxonomic revision of this primate group.Website 
• Nekaris KAI, Arnell AP, Svensson MS, 'Selecting a conservation surrogate species for small fragmented habitats using ecological niche modelling'
  Animals 5 (1) (2015) pp.27-40
  ISSN: 2076-2615 eISSN: 2076-2615
  Abstract Flagship species are traditionally large, charismatic animals used to rally conservation efforts. Accepted flagship definitions suggest they need only fulfil a strategic role, unlike umbrella species that are used to shelter cohabitant taxa. The criteria used to select both flagship and umbrella species may not stand up in the face of dramatic forest loss, where remaining fragments may only contain species that do not suit either set of criteria. The Cinderella species concept covers aesthetically pleasing and overlooked species that fulfil the criteria of flagships or umbrellas. Such species are also more likely to occur in fragmented habitats. We tested Cinderella criteria on mammals in the fragmented forests of the Sri Lankan Wet Zone. We selected taxa that fulfilled both strategic and ecological roles. We created a shortlist of ten species, and from a survey of local perceptions highlighted two finalists. We tested these for umbrella characteristics against the original shortlist, utilizing Maximum Entropy (MaxEnt) modelling, and analysed distribution overlap using ArcGIS. The criteria highlighted Loris tardigradus tardigradus and Prionailurus viverrinus as finalists, with the former having highest flagship potential. We suggest Cinderella species can be effective conservation surrogates especially in habitats where traditional flagship species have been extirpated.Website 
• Musing L, Suzuki K, Nekaris KAI, 'Crossing international borders: the trade of slow lorises, Nycticebus spp., as pets in Japan'
  Asian Primates Journal 5 (1) (2015) pp.12-23
  ISSN: 1979-1631
  Abstract Japan’s involvement in the wildlife trade is widely recognised. We investigated the extent and legitimacy of trade of slow lorises, Nycticebus spp., as pets in Japan. We collected data from online videos, pet shops, and selected informants between May and July 2014 in Japan, and supplemented these with data from CITES Trade Database and confiscation records. We recorded 114 slow lorises in 93 Japanese online videos, and across 20 pet shops we recorded the sale of 74 individuals of six threatened species, including 12 Critically Endangered N. javanicus and two hybrids, each costing USD 3,290-8,650, with animals displayed with falsified CITES permits. From 1985 to 2013, CITES data specify that Japan imported more slow lorises than any other country (n=633), and 400 individuals were confiscated entering Japan, between 2000 and 2013. Penalties imposed on law breakers are weak and our investigations highlight inadequacies in Japan’s enforcement of national law and CITES regulation. To combat the illegal trade and demand for wildlife as pets, we emphasise the need for: stronger penalties,
  improved legislative regulation, provision of educational materials and training programs to border control staff and the public. Finally, we strongly urge the continued monitoring of the slow loris trade in Japan. 
  Website 
• Grow N, Wirdateti, Nekaris KAI, 'Does toxic defence in Nycticebus spp. relate to ectoparasites? The lethal effects of slow loris venom on arthropods'
  Toxicon 95 (2014) pp.1-5
  ISSN: 0041-0101
  Abstract The venom produced by slow lorises (Nycticebus spp.) is toxic both intra- and inter-specifically. In this study we assessed the ecoparasite repellent properties of their venom. We tested venom from two Indonesian slow loris species: Nycticebus javanicus and Nycticebus coucang. Arthropods directly exposed to brachial gland secretions mixed with saliva from both species were immediately impaired or exhibited reduced activity (76%), and often died as a result (61%). We found no significant difference in the result of 60-min trials between N. coucang and N. javanicus [X2(1, n = 140) = 2.110, p = 0.3482]. We found evidence that the degree of lethality of the venom varies according to the arthropod taxa to which it is exposed. While most maggots (84%) were initially impaired from the venom after 10 min, maggots died after a 1 h trial 42% of the time. In contrast, at the end of 1 h trial, spiders died 78% of the time. For all arthropods, the average time to death from exposure was less than 25 min (M = 24.40, SD = 22.60). Ectoparasites including ticks, members of the arachnid order, are known to transmit pathogens to hosts and may be an intended target of the toxic secretions. Our results suggest that one function of slow loris venom is to repel parasites that affect their fitness, and that their topical anointing behaviour may be an adaptive response to ectoparasites.Website 
• Moore RS, Wihermanto, Nekaris KAI, 'Compassionate conservation, rehabilitation, & translocation of Indonesian slow lorises'
  Endangered Species Research 26 (2014) pp.93-102
  ISSN: 1863-5407
  Abstract With the number of threatened species in rescue centres rising, scientific reports on the functioning and success of such centres is essential. Compassionate conservation tries to bridge the gap between animal welfare advocates and conservation biologists, recognising the benefits of preserving a species and its individuals. A case in point is that of Indonesia’s threatened slow lorises Nycticebus spp., where illegal trade is decimating wild populations of these primates. We present 4 yr of data from Ciapus Primate Centre in Indonesia, which received 180 slow lorises between 2008 and 2011. We show that >85% of these primates were unsuitable for reintroduction; 23 slow lorises that were deemed suitable for reintroduction were released to the wild between 2010 and 2013 and were followed with radio tracking. Eleven of them died (on average 76 d post-release), 1 was recaptured (148 d post-release), 6 are no longer being monitored (after, on average, 263 d post-release) and their status is unknown, and 5 are still being monitored (average 226 d post-release, as of December 2013). The challenges posed by work with slow lorises in the Ciapus Primate Centre over these 4 yr, with release success highly variable, show that even with concerted effort, rescue centres need to consider alternative options. We review such options, considering the pros and cons of euthanasia, life in captivity and reintroduction to the wild. We conclude that in today’s global conservation crisis, it can only be beneficial to combine the expertise of animal welfare practitioners and conservation biologists.Website 
• Nekaris KAI, 'Extreme primates: ecology and evolution of Asian lorises'
  Evolutionary Anthropology 23 (2014) pp.177-187
  ISSN: 1060-1538 eISSN: 1520-6505
  Website 
• Nijman V, Nekaris KA, 'Traditions, taboos and trade in slow lorises in Sundanese communities in southern Java, Indonesia'
  Endangered Species Research 25 (1) (2014) pp.79-88
  ISSN: 1863-5407
  Abstract

  For endangered species management it is imperative that there is a shared understanding of the different beliefs, opinions and factual knowledge about different aspects of the conservation programme. To allow this to be taken into account, detailed assessments need to be made of these views, how they differ between individuals, and crucially, what beliefs and views are shared. These assessments require tools from both the natural and the social sciences. Here we focus on the beliefs, opinions and knowledge about Javan slow lorises in 12 Sundanese communities of West Java, Indonesia. Javan slow lorises are small, nocturnal, venomous primates that are threatened by habitat loss and over-exploitation for the illegal pet trade. Based on detailed interviews with 79 informants, and using cluster and cultural domain analyses, we identify and document five different topics, i.e. Trade and exploitation, Taboos against disturbing or capturing, Venomous primates, Blood myths and Mystical powers. The most salient topics were factual (activity pattern, habitat use) and the taboo on collecting or disturbing slow lorises. In communities where there were strong traditions, taboos and beliefs in place, slow lorises were found in seemingly unlikely habitats, in or adjacent to the villages; where these were absent slow lorises were less known to the communities and the slow lorises were either less abundant or absent. We conclude that it is imperative that any management programme or species action plan implemented in West Java has to take into account the traditions, beliefs and taboos regarding slow lorises.

  Website 
• Rode-Margono EJ, Nekaris KAI, 'Impact of climate and moonlight on a venomous mammal, the Javan slow loris (Nycticebus javanicus)'
  Contributions to Zoology 83 (4) (2014) pp.217-225
  ISSN: 1383-4517
  Abstract Predation pressure, food availability, and activity may be affected by level of moonlight and climatic conditions. While many nocturnal mammals reduce activity at high lunar illumination to avoid predators (lunarphobia), most visually-oriented nocturnal primates and birds increase activity in bright nights (lunarphilia) to improve foraging efficiency. Similarly, weather conditions may influence activity level and foraging ability. We examined the response of Javan slow lorises (Nycticebus javanicus Geoffroy, 1812) to moonlight and temperature. We radio-tracked 12 animals in West Java, Indonesia, over 1.5 years, resulting in over 600 hours direct observations. We collected behavioural and environmental data including lunar illumination, number of human observers, and climatic factors, and 185 camera trap nights on potential predators. N. javanicus reduced active behaviours in bright nights. Although this might be interpreted as a predator avoidance strategy, animals remained active when more observers were present. We did not find the same effect of lunar illumination on two potential predators. We detected an interactive effect of minimum temperature and moonlight, e.g. in bright nights slow lorises only reduce activity when it is cold. Slow lorises also were more active in higher humidity and when it was cloudy, whereas potential predators were equally active across conditions. As slow lorises are well-adapted to avoid/defend predators by crypsis, mimicry and the possession of venom, we argue that lunarphobia may be due to prey availability. In bright nights that are cold, the combined effects of high luminosity and low temperature favour reduced activity and even torpor. We conclude that Javan slow lorises are lunarphobic – just as the majority of mammals.Website 
• Nekaris KAI, Moore RS, Rode EJ, Fry BG, 'Mad, bad, and dangerous to know: ecology and evolution of slow loris venom'
  BMC Journal of Venomous Animals and Toxins Including Tropical Diseases 19 (21) (2013)
  ISSN: 1678-9199
  Website 
• Nekaris KA, Starr C, 'Obligate exudativory characterizes the diet of the pygmy slow loris Nycticebus pygmaeus'
  American Journal of Primatology 75 (10) (2013) pp.1054-1061
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract Few primate species are known to excavate plant sources to procure exudates and other foods via active gouging. It is now apparent that slow lorises belong to this rare guild of obligate exudativorous primates. We investigate the diet of the pygmy loris (Nycticebus pygmaeus) in a mixed deciduous forest in the Seima Protection Forest, Eastern Cambodia, and attempted to determine the importance of this resource in their diet. Feeding behaviors of six females and seven males were observed using radio-tracking to facilitate follows, and nine fecal samples were collected in February–May and January–March in 2008 and 2009 respectively. We observed 168 feeding bouts, during which the animals ate exudates (76); fruits (33); arthropods (27); flower parts (21); fungi (3); parts of bamboo culms (7); and reptiles (1). We filmed 19 bouts of exudativory, and observed animals consuming exudates in an orthograde posture, or standing quadrupedally over the exudate source. Pygmy lorises also gouged bamboo to collect lichen and fungi, or broke open dead culms to access invertebrates. Feeding occurred on terminal tree branches (24), tree trunks (21), bamboo (13), the middle of branches (7), and the undergrowth (1). The fecal samples contained plant parts, small-sized arthropods (primarily Coleoptera and Lepidoptera), reptile scales, animal bones, and animal hairs. Pygmy slow lorises are morphologically specialized for processing and digesting exudates, displaying small body sizes, specialized dentitions, elongated, and narrow tongues, large caecums, short duodenums, expanded volar pads, and modified hindlimbs. These features, combined with the prevalence of exudates in their diet across seasons, and ill health when exudates are missing from their diet in captivity, points to this species being an obligate exudativore.Website 
• Nekaris KA-I, Campbell N, Coggins TG, Rode EJ, Nijman V, 'Tickled to Death: Analysing Public Perceptions of ‘Cute’ Videos of Threatened Species (Slow Lorises – Nycticebus spp.) on Web 2.0 Sites'
  PLoS ONE 8 (7) (2013)
  ISSN: 1932-6203
  Abstract

  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.

  Website 
• Munds RA, Nekaris KAI, Ford SM, 'Taxonomy of the Bornean slow loris, with new species Nycticebus kayan'
  American Journal of Primatology 75 (1) (2013) pp.46-56
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract

  More species of nocturnal primates are now recognized than in the past, because many are cryptic species. Subtle morphological disparities, such as pelage pattern and color variation, vocal cues, and genetics have aided in elucidating the number of diagnosable species in a genus. The slow lorises (genus Nycticebus) once included only two species, but recent taxonomic studies resulted in the description of three additional species; further incompletely explored variability characterizes each of the currently described species. The Bornean loris in particular is characterized by pelage and body size variation. In this study, we explored facemask variation in the Bornean loris (N. menagensis). Differing facemask patterns, particularly influenced by the amount of white on the face, significantly clustered together by geographic regions, separated by notable geographic boundaries. Our results support the recognition of four species of Bornean lorises: N. menagensis, N. bancanus, N. borneanus, and N. kayan. Genetic studies are required to support these findings and to refine further our understanding of the marked variability within the Bornean loris populations.

  Website 
• Neilson E, Nijman V, Nekaris KAI, 'Conservation Assessments of Arboreal Mammals in Difficult Terrain: Occupancy Modeling of Pileated Gibbons (Hylobates Pileatus)'
  International Journal of Primatology 34 (2013) pp.823-835
  ISSN: 0164-0291 eISSN: 1573-8604
  Abstract

  Long-term monitoring programs, wildlife surveys, and other research involving species population assessment require reliable data on population status. Given the logistically challenging nature of some species’ habitats and cryptic behaviors, collecting these data can prove to be a considerable barrier. We used detection/nondetection data from pileated gibbons (Hylobates pileatus) in the Cardamom Mountains of southwest Cambodia to estimate their population occupancy and detectability. We modeled occupancy using elevation, tree height, tree density, tree diversity, and disturbance covariates. Modeling demonstrated that 83% of the sites are occupied by Hylobates pileatus and that the detectability of the species varies positively with elevation. No clear relationship between habitat quality covariates and occupancy of Hylobates pileatus emerged. Effort analysis based on model estimates demonstrated that at high elevations, less than half the number of site visits is needed to attain the same detectability estimate precision as across all elevations. We suggest that human activities at low elevations, which affect forest composition, are the central factors impacting the detectability and occupancy of Hylobates pileatus. Longer sampling durations and/or a higher number of site visits, especially at lower elevations, increase precision of the occupancy estimator for the least effort. For effective future monitoring and research for this and similar species, using this relatively simple method, applied with repeat site visits, would allow a longitudinal comparison of detection at sites in difficult terrain.

  Website 
• Nijman V, Nekaris KAI, 'Loud Calls, Startle Behaviour, Social Organisation and Predator Avoidance in Arboreal Langurs (cercopithecidae: Presbytis)'
  Folia Primatologica 83 (2013) pp.274-287
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract

  Linking predator avoidance and social organisation predicts that large groups are favoured under heavy predation pressure but that small, inconspicuous groups may do equally well by avoiding detection altogether. We explored the relationships between antipredator behaviour (vocalisation, concealment, fleeing), detectability (colouration, group size) and social organisation in arboreal langurs. Three clear antipredation conditions emerged: (1) exemplified by Presbytis melalophos--brightly-coloured species (red, yellow) with contrasting colours, living in large unimale-multifemale groups (>10 individuals), that vocalised frequently and that flee loudly through the canopy; (2) exemplified by P. comata--greyish species (some contrasting colours), living in intermediate, unimale-multifemale or 1-male-1-female groups (approx. 7 individuals), that vocalise infrequently and that flee through the middle forest layers; (3) exemplified by P. frontata--dull-coloured species without contrast, living in small unimale-multifemale or 1-male-1-female groups (<5 individuals), that may freeze upon detection, and that may vocalise mainly during the night. Crypsis as an antipredator strategy is restricted to taxa that occur in 1-male-1-female groups. This wide range of antipredator strategies within a monophyletic taxon living in the same general area with a similar suite of predators facing similar predator pressures appears to be unique among the order Primates.

  Website 
• Starr C R, Nekaris K A I, Leung L, 'Hiding from the moonlight: luminosity and temperature affect activity of Asian nocturnal primates in a highly seasonal forest'
  PLoS ONE 7 (4) (2012) pp.e36396-
  ISSN: 1932-6203
  Website 
• Starr C, Nekaris K, Streicher U, Leung L, 'Field surveys of the vulnerable pygmy slow loris Nycticebus pygmaeus using local knowledge in Mondulkiri Province, Cambodia'
  Oryx 45 (2011) pp.135-142
  ISSN: 0030-6053 eISSN: 1365-3008
  Abstract The pygmy slow loris Nycticebus pygmaeus is a little-studied primate endemic to Vietnam, Laos, southern China and eastern Cambodia. Our study aimed to gain local knowledge on the distribution and ecology of, and threats to, the species by interviewing hunters, traders and wildlife protection staff, and to verify this information using a spotlighting survey in three major reserves in Mondulkiri Province, Cambodia. Encounter rates of pygmy loris were assessed along 29 transects (129.5 km), yielding observations of 26 individuals. Mean encounter rates were 0.40 km-1 in Seima Protection Forest, 0.10 km-1 in Phnom Prich Wildlife Sanctuary and 0.00 km-1 in Mondulkiri Protected Forest. Informants had knowledge of where populations occurred, their diet, sociality and habitat preferences. Widespread large population declines were reported and informants linked this to high hunting pressure, particularly in 2001 and 2002. In late 2008 and 2009 we resurveyed three transects that had high encounter rates in early 2008 and failed to detect any lorises. Local informants reported high hunting pressure during the previous wet season in two of these sites, and a gold mine development was underway in the third site. Urgent actions are required to address these population declines and to assess the conservation status of pygmy lorises throughout eastern Cambodia.Website 
• Schwitzer C, Glatt L, Nekaris K, Ganzhorn J, 'Responses of animals to habitat alteration: an overview focussing on primates'
  Endangered Species Research 14 (2011) pp.31-38
  ISSN: 1863-5407
  Abstract Original habitat of animal species is being destroyed at an accelerating rate. This is usually associated with an alteration of the remaining habitat, which becomes degraded and/or fragmented. In many regions, forests are cleared to make way for plantations or other agricultural use, and animal species are forced to coexist with humans. In some countries, forests are in the process of being restored for wildlife. As even long-established and well-protected areas typically comprise mosaics of habitats with different degrees of degradation, the future conservation of many species will depend on the capacity of such altered habitats to support their populations. During the last 15 yr, more and more studies have addressed the way that different species respond to the human-induced change of their habitats. These responses are varied, and range from population decline to adaptation and development of new behavioural strategies. Whereas some species rely heavily on intact primary forests, others can adapt to secondary forests and forest-agriculture mosaics. Habitat change has been shown to affect many aspects of the ecology and behaviour of animals. Changes in dietary composition and diversity, population density, group size and adult sex ratio in groups are some examples. This Theme Section of Endangered Species Research collates a number of case studies on how animals, and particularly primates, respond to the alteration of their habitat.Website 
• Nijman V, Nekaris K, Donati G, Bruford M, Fa J, 'Primate conservation: measuring and mitigating trade in primates'
  Endangered Species Research 13 (2) (2011) pp.159-161
  ISSN: 1863-5407
  Abstract

  Trade in primates is seen as a significant impediment to their conservation. Primates are traded both domestically and internationally, in order to supply, amongst others, biomedical industries and pharmaceutical markets, the entertainment business, or pet markets. Primate meat is consumed globally, whereas body parts are used as ingredients in traditional medicine or sold as curios. All international trade in primates is regulated through the Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES), to which all but 2 primate range countries are signatory. The last 15 years has seen a linear increase in the export of live primates (each year 3500 more individuals are exported), with China being, numerically, the largest exporter. While the trade in live primates worldwide involves tens, if not hundreds, of thousands of individuals a year, the trade in dead primates involves millions of animals a year. We introduce here a series of studies dealing with various aspects of the primate trade. We hope that these studies will urge others to quantify the extent of trade in primates alive and dead in both domestic and international contexts, allowing us to find ways to mitigate the consequences of this trade to the conservation of primates.

  Website 
• Nekaris K, Nijman V, Godfrey L, 'Anthropology: follow field primatologists'
  Nature 471 (488) (2011)
  ISSN: 0028-0836
  Abstract Field primatologists search the archaeological record of tool-using primates to gain insight into their cultures and traditions. Similarly, researchers of primate communication have set up a linguistic framework to investigate its intricacies in the context of the evolution of human language and music.Website 
• Rode E, Nekaris K, Schwitzer C, Hoffman M, 'Mirza Zaza - preliminary conservation status assessment for the Data Deficient northern giant mouse lemur'
  Lemur News (2011) pp.11-12
  ISSN: 1608-1439
  Abstract No abstract available
• Svensson MS, Samudio R, Bearder SK, Nekaris KAI, 'Density estimates of Panamanian owl monkeys (Aotus zonalis) in three habitat types'
  American Journal of Primatology 72 (2) (2010) pp.187-192
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract The resolution of the ambiguity surrounding the taxonomy of Aotus means data on newly classified species are urgently needed for conservation efforts. We conducted a study on the Panamanian owl monkey (Aotus zonalis) between May and July 2008 at three localities in Chagres National Park, located east of the Panama Canal, using the line transect method to quantify abundance and distribution. Vegetation surveys were also conducted to provide a baseline quantification of the three habitat types. We observed 33 individuals within 16 groups in two out of the three sites. Population density was highest in Campo Chagres with 19.7 individuals/km2 and intermediate densities of 14.3 individuals/km2 were observed at Cerro Azul. In la Llana A. zonalis was not found to be present. The presence of A. zonalis in Chagres National Park, albeit at seemingly low abundance, is encouraging. A longer‐term study will be necessary to validate the further abundance estimates gained in this pilot study in order to make conservation policy decisions.Website 
• Nekaris K, Shepherd C, Starr C, Nijman V, 'Exploring cultural drivers for wildlife trade via an ethnoprimatological approach: a case study of slender and slow lorises (Loris and Nycticebus) in South and Southeast Asia'
  American Journal of Primatology 72 (10) (2010) pp.877-886
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract Illegal and unsustainable trade in wildlife is a major conservation challenge. For Asian primates, economic and cultural traditions, and increased forest access mean that trade may have become detrimental for certain species. Slow and slender lorises (Nycticebus and Loris) are primates particularly prevalent in trade, determined until now by focused counts of lorises in regional markets. Here, we use international trade statistics and a participant-observer approach to assess culturally specific drivers for trade in lorises in South and Southeast Asia, to provide a broader context to help mitigate this practice. Analysis of international records for the last 30 years revealed that live animal trade was more prevalent than trade in body parts (slow lorises, 86.4%; slender lorises, 91.4%), with Laos, Cambodia, and Thailand the largest exporters. We then examine drivers of international and domestic trade based on long-term data from 1994-2009 in Sri Lanka, Cambodia, and Indonesia. We show that slender lorises are important in Sri Lankan folklore, but their use as pets and for traditional medicine is rare. Trade in Bengal slow and pygmy lorises in Cambodia for use in traditional medicines, a practice with deeply historical roots, is widespread. Despite its own set of myths about the magical and curative properties of lorises, trade in Javan, Bornean, and greater slow lorises in Indonesia is largely for pets. Conservation practices in Asia are often generalized and linked with the region's major religions and economies. We show here that, in the case of wildlife trade, culturally specific patterns are evident among different ethnic groups, even within a country. Revealing such patterns is the foundation for developing conservation management plans for each species. We suggest some participatory methods for each country that may aid in this process.Website 
• Nijman V, Nekaris K, 'Effects of deforestation on attitudes and levels of tolerance towards commensal primates (Cercopithecidae) in Sri Lanka'
  International Journal of Pest Management 56 (2) (2010) pp.153-158
  ISSN: 0967-0874 eISSN: 1366-5863
  Abstract Attitudes of people to wildlife, particularly to animals that live in close proximity to them, are an important element of conservation efforts and management. Attitudes may vary according to age and levels of conflict. We assessed the influence of proximity of forest on the attitudes of people towards two commensal primates, the purple-faced langur and the toque macaque. Data were collected in Sri Lanka by interviews in three villages where there is no continuous forest remaining and in three villages with adjacent forest. We found high levels of tolerance towards commensal primates, but significantly higher levels of negative perceptions in villages where forest was no longer present. Perceptions were not related to age or sex. The total disappearance of forest, with primates being dependent on fruit crops and living permanently on the village grounds, inevitably leads to conflict. These changing views have important management implications. Animals surviving in a human-dominated landscape may become more common, and the experiences in Sri Lanka may provide insight into what the future holds for other sites.Website 
• Starr C, Nekaris K, Streicher U, Leung L, 'Traditional use of slow lorises Nycticebus bengalensis and N. pygmaeus in Cambodia: an impediment to their conservation'
  Endangered Species Research 12 (1) (2010) pp.17-23
  ISSN: 1863-5407
  Abstract In Cambodia, periodic reports since the 1990s have identified the sale of large numbers of dried pygmy Nycticebus pygmaeus and northern slow lorises N. bengalensis in traditional medicine stores. We used interviews and questionnaires to identify the uses and users of lorises, elucidate factors affecting selection of loris medicines, and determine whether access to alternative therapies may reduce the use of loris medicines. Pygmy lorises were found to be the most commonly requested animal from traditional medicine stores in Cambodia" s capital Phnom Penh, and the primary users recalled by sellers were women between the ages of 25 and 45 from middle to upper class backgrounds. Slow lorises were predominantly used in a tonic for women after childbirth, stomach problems, healing wounds and broken bones, and in the treatment of sexually transmitted diseases. Lorises were reported to be sourced from provinces with large protected areas within their distribution ranges. Market values of both species more than doubled from 1997 to 2007; however, the majority of respondents expressed reluctance to substitute loris medicines with alternatives, indicating that promotion of alternatives would be an inappropriate conservation tool. Education and enhanced law enforcement are vital to conserve slow lorises in Cambodia. Website 
• Nijman V, Nekaris K, 'Testing a model for predicting primate crop-raiding using crop- and farm-specific risk values'
  Applied Animal Behaviour Science 127 (41002) (2010) pp.125-129
  ISSN: 0168-1591
  Abstract Crop-raiding by primates is increasingly known to cause conflict between humans and primates, and due to their opportunism, adaptability, intelligence and manipulative abilities, primates can be significant agricultural pests. Levels of crop-raiding are dependent on time of year, crop type, size and location of the farm, and primate species involved, making it difficult for farmers to predict susceptibility to crop-raiding accurately. We use a simple method for calculating the likelihood of crop damage by primates using crop susceptibility to predict the frequency of crop damage for individual farms. The method relies on calculating incidence rates of crop-raiding for individual crops using pooled data from all farms in the sample, and summing these rates as to reach a farm's risk value (RV) to primate crop-raiding. From 273 farms in southwestern Sri Lanka data were collected on crop-raiding by two species of primate, the arboreal folivorous purple-faced langur and the terrestrial frugivorous toque macaque. Data from 93 farms were used to calculate crop-raiding incidence rates for seven commonly grown crops, and we tested the applicability of the model using the remainder of the dataset. Incidence rates of raiding for crops differed for the two species of primate, albeit not in a uniform manner. Farms appear to be more susceptible to crop-raiding by langurs than by macaques, with higher RVs for langurs than for macaques: this is not related to the behaviour of the farmer as for both species four-fifth of the farmers that experience crop-raiding actively chase primates away. Our model using RVs works well for predicting crop-raiding in langurs as crop-raided farms have significantly higher RVs. It works less well for macaques, which may be related to their terrestriallity allowing them to range over larger areas and raid farms opportunistically.Website 
• Moore R, Nekaris K, Eschmann C, 'Habitat use by western purple-faced langurs Trachypithecus vetulus nestor (Colobinae) in a fragmented suburban landscape'
  Endangered Species Research 12 (3) (2010) pp.227-234
  ISSN: 1863-5407
  Abstract As natural habitats around the globe disappear, humans and non-human primates become increasingly engaged in complex interactions, both peaceful and hostile. Sri Lanka" s endemic western purple-faced langur Trachypithecus vetulus nestor persists in the majority of its range in complete sympatry with humans. Listed as Critically Endangered by the IUCN and one of the world" s top 25 most endangered primates, their survival appears dependent on the tolerance of humans with whom they coexist. Our aims were to augment the limited information on T. v. nestor focusing on group composition and behavioural adaptations in relation to its heavily fragmented habitat. Research was carried out in Talangama Wetlands, Sri Lanka in 2007. The 5 groups varied in size from 11 to 19 individuals Monkeys regularly used fences, rooftops, power lines, and agricultural trees. Two groups of 19 and 16 individuals had home ranges of 3.06 and 2.86 ha respectively. They consumed a variety of foods including fruits, showing dietary flexibility; the fruits and leaves of Arctocarpus heterophyllus were the langurs" most frequently consumed foods. The langurs seem to be adapting to these human-modified environments. Notwithstanding, areas for concern include potentially fatal dangers when crossing between fragments, increasing human-primate conflict and inter-group conflict, and permanent genetic isolation. Website 
• Nekaris K, Llano Sanchez K, Thorn J, Winarti I, Nijman V, 'Javan Slow Loris (Nycticebus Javanicas)'
  (2009) pp.44-46
  Abstract No abstract available
• Thorn J S, Smith D, Nijman V, Nekaris K A I, 'Ecological niche modelling as a technique for assessing threats and setting conservation priorities for Asian slow lorises (Primates: Nycticebus)'
  Diversity and Distributions 15 (2) (2009) pp.289-298
  ISSN: 1366-9516
  Abstract

  Aim: Data on geographical ranges are essential when defining the conservation status of a species, and in evaluating levels of human disturbance. Where locality data are deficient, presence-only ecological niche modelling (ENM) can provide insights into a species' potential distribution, and can aid in conservation planning. Presence-only ENM is especially important for rare, cryptic and nocturnal species, where absence is difficult to define. Here we applied ENM to carry out an anthropogenic risk assessment and set conservation priorities for three threatened species of Asian slow loris (Primates: Nycticebus). Location: Borneo, Java and Sumatra, Southeast Asia. Methods: Distribution models were built using maximum entropy (MaxEnt) ENM. We input 20 environmental variables comprising temperature, precipitation and altitude, along with species locality data. We clipped predicted distributions to forest cover and altitudinal data to generate remnant distributions. These were then applied to protected area (PA) and human land-use data, using specific criteria to define low-, medium- or high-risk areas. These data were analysed to pinpoint priority study sites, suitable reintroduction zones and protected area extensions. Results: A jackknife validation method indicated highly significant models for all three species with small sample sizes (n = 10 to 23 occurrences). The distribution models represented high habitat suitability within each species' geographical range. High-risk areas were most prevalent for the Javan slow loris (Nycticebus javanicus) on Java, with the highest proportion of low-risk areas for the Bornean slow loris (N. menagensis) on Borneo. Eighteen PA extensions and 23 priority survey sites were identified across the study region. Main conclusions: Discriminating areas of high habitat suitability lays the foundations for planning field studies and conservation initiatives. This study highlights potential reintroduction zones that will minimize anthropogenic threats to animals that are released. These data reiterate the conclusion of previous research, showing MaxEnt is a viable technique for modelling species distributions with small sample sizes.

  Website 
• Biebouw K, Bearder S, Nekaris K, 'Tree Hole Utilisation by the Hairy-Eared Dwarf Lemur (Allocebus trichotis) in Analamazaotra Special Reserve'
  Folia Primatologica 80 (2) (2009) pp.89-103
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract

  In this study we describe tree hole characteristics and use by the hairy-eared dwarf lemur (Allocebus trichotis) to determine habitat needs, potential functions of tree holes and sleeping group composition. We radio-tracked 6 adult individuals between April and November 2007 in the Analamazaotra Special Reserve. Tree holes were 1-9 m high (median: 7 m), in living trees measuring 26-54 cm in diameter at breast height (median: 32 cm), and could be a limiting resource. Each individual used 4 or 5 tree holes and had high nest fidelity. Animals most often slept socially in mixed-sex groups of 2-6 individuals and occasionally shared a tree hole with white-tailed tree rats (Brachytarsomys albicauda). We identified two sleeping groups: one composed of 2 adult males, 2 adult females and 2 juveniles; one composed of at least 2 adult females and 2 juveniles. Although tree holes were generally group exclusive, some intergroup sleeping was observed. Tree holes could have antipredator and thermoregulatory functions. Further research into sleeping hole availability, nest use and the degree of niche separation or competition between sympatric Cheirogaleidae and other tree hole users (e.g. endemic rodents) is needed to assess better the conservation needs of these species.

  Website 
• Nekaris K, Nijman V, 'Attention for Sri Lanken monkey paints a bleak picture yet gives a glimmer of hope'
  Oryx 42 (2008) pp.487-488
  ISSN: 0030-6053 eISSN: 1365-3008
  Abstract No abstract availableWebsite 
• Smith D, Thompson S, Nekaris K, 'An assessment of conservation initiatives and priorities, and the habitat risks of human land use to the primates (colobinae, hylobatidae and pongidae) of Kalimantan'
  Folia Primatologica 79 (5) (2008) pp.385-386
  ISSN: 0015-5713 eISSN: 1421-9980
  
• Parker L, Nijman V, Nekaris K, 'When there is no forest left: fragmentation, local extinction, and small population sizes in the Sri Lankan western purple-faced langur'
  Endangered Species Research 5 (1) (2008) pp.29-36
  ISSN: 1863-5407
  Abstract The western purple-faced langur Trachypithecus vetulus nestor is a Critically Endangered primate endemic to Sri Lanka according to the International Union for Conservation of Nature (IUCN). Its population decline is inferred mainly due to vast habitat loss. Few recent data are available regarding its distribution or abundance. The aim of our study was to assess the conservation status of T. v. nestor throughout its known historical range by establishing presence/absence and correlating these data with semi-structured interviews determining human perceptions and threats. Twenty-six sites were investigated, with presence of T. v. nestor being confirmed at 11 and indicated through questionnaires only at 2 additional sites. The present distribution of T. v. nestor is severely fragmented both locally and regionally. More than half of the populations comprised 1 or 2 groupsonly and, being isolated in a matrix of urban landscapes, are close to unviable. Conflicts between local people and T. v. nestor were identified at such sites, where reports of troops crop-raiding gardensand plantations were associated with negative perceptions. Consequences for crop-raiding langurs ranged from shouting to (infrequently) shooting and killing by dogs. Negative views towards T.v. nestor were significantly related to the length of time they spent in anthropogenic landscapes, while more positive views were significantly associated with the length of time T. v. nestor spent in forested areas. Evidence of local extinctions means that the remaining, yet increasingly fragmented habitat of the highly arboreal T. v. nestor is critical for survival. Intervention to manage this human-wildlife conflict is vital. An integrated approach using international and local conservation authorities is highly recommended.Website 
• Nekaris K A I, Blackman G, Nijman V, 'Conservation implications of low encounter rates of five nocturnal primate species (Nycticebus spp.) in Asia'
  Biodiversity and Conservation 17 (4) (2008) pp.733-747
  ISSN: 0960-3115 eISSN: 1572-9710
  Abstract 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.Website 
• Eschmann C, Moore R, Nekaris K, 'Calling patterns of Western purple-faced langurs (Mammalia: Primates: Cercopithecidea: Trachypithecus vetulus nestor) in a degraded human landscape in Sri Lanka'
  Contributions to Zoology 77 (2) (2008) pp.57-65
  ISSN: 1383-4517
  Abstract The study of calling patterns is a useful non-invasive method for determining population densities and the taxonomic relationships of rare or cryptic animal species. The Western purple-faced langur Trachypithecus vetulus nestor, endemic to Sri Lanka" s lowland rainforests, is severely impacted by forest fragmentation, with most remaining populations living almost completely in home gardens. Due to their shy nature, little is known about the behaviour of this subspecies; analysing the regular loud calls emitted by these langurs could allow for improvement of census techniques, clarification of their taxonomy, and an understanding of the impact of forest destruction on their behaviour. In 2007, we recorded the calling patterns of five male T. v. nestor at Talangama Wetlands. Time, duration, weather conditions, and stimulant of 253 calls were noted. Loud calls comprised three structural units: harsh barks, whoops and residuals. The average call contained 4 phrases and 3.8 residuals, was 38 seconds in length, had an average maximum frequency of 3.5 kHz, a formant frequency of 0.36 kHz, and a fundamental frequency of 0.2 kHz. Significant differences were found between individuals for the number of phrases and residuals within a call, two different phrase lengths, the formant frequency and the fundamental frequency. The earliest call occurred at 05:27 hrs, while the latest was made at 17:57 hrs. The greatest percentage of calls (73.5%) was heard in the morning (05:00-09:59 hrs), mostly stimulated by territorial battles with neighbouring troops. These results show that vocalisations can be used to distinguish individual males; as langurs are more often heard than seen, and most troops contain only a single adult male, vocalisations may be used to determine the number of troops in an area. Calls of this taxon also differed from the other subspecies, suggesting that they may be used to distinguish subspecies and their boundaries. Finally, calling behaviour differed from other subspecies. Deforestation may be a direct cause of different calling patterns. These baseline data form a valuable starting point for further studies of this Critically Endangered primate.
• Aldrich B, Molleson L, Nekaris K, 'Vocalizations as a conservation tool: an auditory survey of the Andean titi monkey Callicebus oenanthe Thomas, 1924 (Mammalia: Primates: Pitheciidae) at Tarangue, Northern Peru'
  Contributions to Zoology 77 (1) (2008) pp.1-6
  ISSN: 1383-4517
  Abstract Titi monkeys (Callicebus), morphologically cryptic primates, have been difficult to survey using traditional sighting-based line transect methods. Callicebus-species regularly engage in loud, ritualized singing bouts, which could allow for the use of alternate, potentially more accurate call-based survey methods to monitor populations. The Andean titi monkey, C. oenanthe, is endemic to a small region of northern Peru, an area subject to widespread and rapid deforestation and human colonization. We conducted a call-based survey of C. oenanthe at Tarangue, a 74 ha private reserve near Moyobamba. Triangulation of calls was used to map groups of titi monkeys on and around the reserve. 73 mapped calls were used to estimate the presence of between three and six groups per listening area - a total of 23 groups entirely or partially within the borders of Tarangue, yielding an estimated population density of 1.41 individuals per ha. Observations were much greater than those resulting from a visually-based survey conducted at Tarangue three years earlier. These higher estimates are probably not only due to this more suitable survey method; incessant destruction of habitat occurring in the area surrounding Tarangue may have caused the reserve to become a refuge for displaced individuals, with diminished opportunities for dispersal and establishment of new territories. Immediate measures to prevent further fragmentation within the Andean titi monkey" s geographic range are essential in order to allow the species to persist. We recommend the use of triangulation of calls for future surveys of titi monkeys.
• Nekaris K, Nijman V, 'CITES Proposal Highlights Rarity of Asian Nocturnal Primates (Lorisidae: Nycticebus )'
  Logo Folia Primatologica 78 (2007) pp.211-214
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract No abstract availableWebsite 
• Nekaris K, 'Horton Plains Slender Loris, Ceylon Mountain Slender Loris'
  (2007) pp.12-13
  Abstract No abstract available
• Nekaris K, Jaffe S, 'Unexpected diversity of slow lorises (Nycticebus spp.) within the Javan pet trade: implications for slow loris taxonomy'
  Contributions to Zoology 76 (3) (2007) pp.187-196
  ISSN: 1383-4517
  Abstract Since the 1950s, Sundaland (Borneo, Java, Sumatra and their surrounding islands) was thought to be inhabited by a single slow loris species, the greater slow loris Nycticebus coucang. Early taxonomies as well as recent morphological and genetic studies, however, point to at least three species native to this region: N. coucang, N. menagensis, and N. javanicus. In the light of this taxonomy, all Sundaland slow lorises, previously considered Least Threatened, have been listed as Vulnerable or Endangered. Of particular concern is the fact that slow lorises are the most common protected primate species in the rampant Southeast Asian pet trade, resulting in their recent transferral to CITES Appendix I precluding all international commercial trade. Due to lack of knowledge regarding morphological differences between the three species, they are still managed as one, with potential serious affects to wild populations, as hard-release of individuals of unknown geographic origin is common. This paper examines morphological variability of 34 live slow lorises, all of which were rescued from the wildlife trade in Java, Indonesia. Morphometric data and diagnostic images were collected, various species descriptions were considered and statistical analyses were conducted and compared with other taxonomists" classifications. A discriminant function analysis provided support for four distinct groupings: Nycticebus coucang and N. javanicus, as well as evidence for two new taxa that correspond closely to N. hilleri and N. ornatus. The morphological traits that varied signifi cantly and the external characteristic trends described in this study that contributed to these groupings might provide a baseline to classify Nycticebus taxa. This information is pertinent for appropriate captive management and specifi c designation of rescued individuals and for designing proper in-situ and ex-situ conservation strategies.
• Nekaris K, Stephens N, 'Not all lorises are slow: rapid arboreal locomotion in Loris tardigradus of Southwestern Sri Lanka'
  American Journal of Primatology 69 (1) (2007) pp.113-121
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract The unique slow-climbing quadrupedalism of Asian lorises has been the subject of numerous studies; however, qualitative observations of more rapid locomotion have occasionally been reported. Field studies of the red slender loris have revealed the habitual use of unexpectedly high-speed locomotion by the so-called -œsloth of the primate world.- Novel video footage permitted the first quantitative kinematic analysis of rapid quadrupedalism in wild lorises. Observations revealed that this previously unexplored behavior is far from infrequent, with 26% of red slender loris locomotor activity being dedicated to high-velocity arboreal quadrupedalism. This locomotor pattern may represent a primitive retention of the rapid, scrambling quadrupedalism that is observed in other strepsirhines, or it may constitute a more recent specialization of this smallest loris taxon.Website 
• Nekaris K, 'Social lives of adult Mysore slender lorises (Loris lydekkerianus lydekkerianus)'
  American Journal of Primatology 68 (12) (2006) pp.1171-1182
  ISSN: 0275-2565 eISSN: 1098-2345
  Abstract Despite the persistent use of the word 'solitary' to describe nocturnal primate social behavior, increasing numbers of studies are revealing sophisticated levels of social interactions among nocturnal primates. This study explores the relationships among 11 adult Mysore slender lorises (Loris lydekkerianus lydekkerianus) studied over 101/2 months in Tamil Nadu, India. When all observations regarding dependent offspring are excluded, the animals spent on average 38% of their activity in various forms of neutral, affiliative, and agonistic behaviors. Affiliative behaviors were the most common type of social interaction, and males in general were more social than females. Low values for Cole's index (CI) of association emphasize that females rarely interacted with same-sex conspecifics, but commonly interacted with males. In turn, males also formed strong affiliative relationships with other adult males. This index also indicates that levels of affiliation are strongest among animals that share sleeping sites. The Hinde index (HI) suggests that males control proximity to females more than the reverse. A female's tolerance of multiple males in her home range and at a sleeping site may be related to high spatial variability of food resources. Such resources may constrain females with costly reproductive strategies (up to two sets of twins per annum) to a small home range. With their larger home ranges, males may be able to monopolize females by initiating social interactions, and also provide a benefit to females by contributing to parental care.Website 
• Buckley C, Nekaris K, Husson S, 'Survey of Hylobates agilis albibarbis in a logged peat-swamp forest: Sabangau catchment, Central Kalimantan'
  Primates 47 (4) (2006) pp.327-335
  ISSN: 0032-8332
  Abstract Few data are available on gibbon populations in peat-swamp forest. In order to assess the importance of this habitat for gibbon conservation, a population of Hylobates agilis albibarbis was surveyed in the Sabangau peat-swamp forest, Central Kalimantan, Indonesia. This is an area of about 5,500 km2 of selectively logged peat-swamp forest, which was formally gazetted as a national park during 2005. The study was conducted during June and July 2004 using auditory sampling methods. Five sample areas were selected and each was surveyed for four consecutive days by three teams of researchers at designated listening posts. Researchers recorded compass bearings of, and estimated distances to, singing groups. Nineteen groups were located. Population density is estimated to be 2.16 (±0.46) groups/km2. Sightings occurring either at the listening posts or that were obtained by tracking in on calling groups yielded a mean group size of 3.4 individuals, hence individual gibbon density is estimated to be 7.4 (±1.59) individuals/km2. The density estimates fall at the mid-range of those calculated for other gibbon populations, thus suggesting that peat-swamp forest is an important habitat for gibbon conservation in Borneo. A tentative extrapolation of results suggests a potential gibbon population size of 19,000 individuals within the mixed-swamp forest habitat sub-type in the Sabangau. This represents one of the largest remaining continuous populations of Bornean agile gibbons. The designation of the Sabangau forest as a national park will hopefully address the problem of illegal logging and hunting in the region. Further studies should note any difference in gibbon density post protection.Website 
• Nekaris K, 'Foraging behaviour of the slender loris: implications for theories of primate origins'
  Journal of Human Evolution 49 (3) (2005) pp.289-300
  ISSN: 0047-2484
  Abstract Members of the Order Primates are characterised by a wide overlap of visual fields or optic convergence. It has been proposed that exploitation of either insects or angiosperm products in the terminal branches of trees, and the corresponding complex, three-dimensional environment associated with these foraging strategies, account for visual convergence. Although slender lorises (Loris sp.) are the most visually convergent of all the primates, very little is known about their feeding ecology. This study, carried out over 10 ½ months in South India, examines the feeding behaviour of L. lydekkerianus lydekkerianus in relation to hypotheses regarding visual predation of insects. Of 1238 feeding observations, 96% were of animal prey. Lorises showed an equal and overwhelming preference for terminal and middle branch feeding, using the undergrowth and trunk rarely. The type of prey caught on terminal branches (Lepidoptera, Odonata, Homoptera) differed significantly from those caught on middle branches (Hymenoptera, Coleoptera). A two-handed catch accompanied by bipedal postures was used almost exclusively on terminal branches where mobile prey was caught, whereas the more common capture technique of one-handed grab was used more often on sturdy middle branches to obtain slow moving prey. Although prey was detected with senses other than vision, vision was the key sense used upon the final strike. This study strongly supports the notion that hunting for animal prey was a key ecological determinant in selecting for visual convergence early on in primate evolution. The extreme specialisations of slender lorises, however, suggest that early primates were not dedicated faunivores and lend further support to the emerging view that both insects and fruits were probably important components of the diet of basal primates, and that exploitation of fruits may account for other key primate traits.Website 
• Nekaris K, Jayewardene J, 'Survey of the slender loris (Primates, Lorisidae Gray, 1821: Loris tardigradus Linnaeus, 1758 and Loris lydekkerianus Cabrera, 1908) in Sri Lanka'
  Journal of Zoology 262 (4) (2004) pp.327-338
  ISSN: 0952-8369
  Abstract In 2001 and 2002, surveys of slender lorises were carried out in Sri Lanka, providing the first recent information on four taxa (Loris lydekkerianus nordicus, L. l. grandis, L. tardigradus tardigradus, and L. t. nycticeboides) endemic to the island. Thirty-one sites across five ecological zones were surveyed. Approximately 766 km were covered in 17 areas where no lorises were found; 192 km were walked or motored in 14 sites yielding 185 sightings of Loris: L. l. nordicus (n=111), L. t. tardigradus (n=69), L. l. grandis (n=4), and L. t. nycticeboides (n=1). Density estimates, based on sightings of animals/km, were: L. t. tardigradus (0.86-13 animals/km) and L. l. nordicus (0.33-50 animals/km). Significantly fewer sightings occurred within protected areas than were made outside of them. Animal densities varied across habitat type with the highest density of lorises occurring in the dry zone in monsoon forests. Presence of Loris is positively associated with insect presence, and negatively associated with primary forest with little undergrowth; taxa differ in their ability to thrive on the edge of human habitations. Human-induced threats include habitat loss, electrocution on live wires, road accidents, the pet trade, and use in traditional medicine. Further behavioural and ecological studies are needed to estimate the habitat requirements for the different taxa of slender loris.Website 
• Nekaris K, 'Observations of Mating, Birthing and Parental Behaviour in Three Subspecies of Slender Loris (Loris tardigradus and Loris lydekkerianus ) in India and Sri Lanka'
  Folia Primatologica 74 (2003) pp.312-336
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract Studies of the life history parameters of slender lorises in captivity have led to conflicting results regarding gestation length, birth seasonality, interspecies variation in litter size and the degree of parental care given to offspring. During the course of field studies of Loris lydekkerianus lydekkerianus, L. l. nordicus and L. tardigradus tardigradus, data were collected on these life history variables, as well as on behaviours relating to mating. All 3 taxa displayed courtship behaviour involving the pursuit of a female by multiple males. Mating corresponded closely with captive observations, with a period of prolonged single intromissions lasting 3-11 min. One gestation period of 163 days was calculated for L. l. lydekkerianus. Births for all 3 taxa were distributed throughout the year, and males were seenmating throughout the year. All 3 taxa gave birth to singletons and twins; no subspecific pattern in litter size was evident. Females carried infants for the first 4 weeks of life and were regularly attended by males, which groomed both the mother and her offspring. After infants had been parked, female L. l. lydekkerianus and L. l. nordicus rarely returned before dawn, though males visited and played with infants. Female L. t. tardigradus maintained proximity with their infants, whilst males were not observed in proximity to infants during the night. All 3 taxa slept in social groups. High-energy milk, in combination with male care, may aid in the potentially high reproductive output of 4 infants per year.Website 
• Nekaris K, Rasmussen D, 'Diet and Feeding Behavior of Mysore Slender Lorises'
  International Journal of Primatology 24 (1) (2003) pp.33-46
  ISSN: 0164-0291 eISSN: 1573-8604
  Abstract We studied the feeding ecology of the Mysore slender loris (Loris lydekkerianus lydekkerianus) for 10.5 mo in a dry scrub forest at Ayyalur Interface Forestry Division, Tamil Nadu, South India. We recorded and analyzed 1240 feeding incidents, which indicate that the lorises were almost exclusively faunivorous, with 96% of all feeding events representing animal prey. Of prey items that could be identified (n = 605), 62.9% were ants and termites. Lorises fed on ge9 orders and 17 families of insects, plus spiders, molluscs, and small vertebrates. Lorises infrequently fed on gums and a legume pod. They usually grabbed prey with one hand, while other appendages firmly held the substrate. Many of the identifiable prey items belong to insect taxa likely to contain toxic chemicals. Consumption of insects inferred to be toxic was accompanied by an elaborate behavioral repertoire of sneezing, slobbering and urine-washing. A high proportion of insects eaten by slender lorises (71%) occurred in patches or aggregations. The utilization of aggregated social insects may have implications for understanding the unusually high degree of gregarious behavior exhibited by the lorises.Website 
• Nekaris K, 'Spacing system of the Mysore slender loris (Loris lydekkerianus lydekkerianus)'
  American Journal of Physical Anthropology 121 (1) (2003) pp.86-96
  ISSN: 0002-9483 eISSN: 1096-8644
  Abstract Loris lydekkerianus lydekkerianus has been shown to have a promiscuous copulatory pattern, to maintain social networks via frequent loud calls, to interact socially throughout the night with all age classes, and to sleep socially. Though these behaviors point towards a multimale social system, no study of their spacing system has yet been provided to support this view. From October 1997-August 1998, I conducted a study of the Mysore slender loris in Ayyalur, India. During 1,400 field hours, data were collected on range use of 3 adult females, 3 adult males, 1 subadult female, and 1 subadult male. Lorises slept in groups averaging 4 individuals, composed of an adult female, her offspring, and 1-2 adult and subadult males. Sleeping sites for three groups were located within 1.9 ha in the center of the study area. The minimum convex polygon in hectares encompassing each animal" s range was determined, as well as overlap among home ranges of individual lorises. Average home range sizes were: adult males, 3.6 ha 0.09; subadult/smaller males, 1.17 ha 0.26; and adult and subadult females, 1.59 ha 0.24. Male ranges overlapped with at least 2-3 other adult males (0.72 ha 0.23). Female ranges overlapped slightly with at least 2 other female ranges (0.22 ha 0.25). Male ranges overlapped those of at least 3 females (0.82 ha 0.51). Patterns of home range and sleeping site support previous suggestions of a multimale social system, similar to aye ayes and some galagos. Website 
• Gursky S, Nekaris K, 'An Introduction to Mating, Birthing and Rearing Systems of Nocturnal Prosimians'
  Folia Primatologica 74 (2003) pp.241-245
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract No abstract availableWebsite 
• Nekaris K, Jayewardene J, 'Pilot Study and Conservation Status of the Slender Loris (Loris tardigradus and L. lydekkerianus) in Sri Lanka'
  Primate Conservation 19 (2003) pp.83-90
  ISSN: 0898-6207
  Abstract No abstract available
• Nekaris K, 'Activity Budget and Positional Behavior of the Mysore Slender Loris (Loris tardigradus lydekkerianus): Implications for Slow Climbing Locomotion'
  Folia Primatologica 72 (4) (2001) pp.228-241
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract Both predator defense and feeding ecology models have been proposed to explain the relatively slow climbing locomotion of the Lorisinae. During a study of the socioecology of the Mysore slender loris (Loris tardigradus lydekkerianus) in Tamil Nadu, India, six categories of behavior and eleven different postures were recorded to estimate a general activity budget for the slender loris, and are examined here particularly in relation to slow climbing locomotor strategies. Reactions to potential predators are also described. The main study population was composed of 15 animals. Activity budgets were compiled in three ways: all instantaneous point samples collected over 1,173 h pooled (n = 13,717), the means of individual lorises (n = 15) and behavior at the moment of first contact (n = 357). No significant difference was found between these three data sets. Approximately 45% of the activity budget was spent in inactive behaviors including sitting vigilant, resting and sleeping. Foraging and traveling comprised nearly half the activity budget, with the rest of the time spent grooming. The most common postures assumed by lorises were sitting and quadrupedal walking. Individual lorises were relatively gregarious and spent up to half their activity budget with other animals. Unlike pottos and angwantibos, lorises did not freeze, head butt or drop from branches in reaction to potential predators, but either ignored them, fled or made loud calls. Cryptic and slow climbing locomotion were used before traveling on open ground between discontinuous substrates, thereby supporting hypotheses relating to predator pressure, and also before capturing fast moving insect prey, supporting hypotheses relating to diet. It is proposed that a divergence in foraging strategies between bushbabies and lorisines may be the best adaptive explanation for their behavioral and morphological differences, including predator defense mechanisms.Website 
• Rasmussen D, Nekaris K, 'Evolutionary History of Lorisiform Primates'
  Folia Primatologica 69 (Supplement 1) (1998) pp.250-285
  ISSN: 0015-5713 eISSN: 1421-9980
  Abstract

  We integrate information from the fossil record, morphology, behavior and molecular studies to provide a current overview of lorisoid evolution. Several Eocene prosimians of the northern continents, including both omomyids and adapoids, have been suggested as possible lorisoid ancestors, but these cannot be substantiated as true strepsirhines. A small-bodied primate, Anchomomys, of the middle Eocene of Europe may be the best candidate among putative adapoids for status as a true strepsirhine. Recent finds of Eocene primates in Africa have revealed new prosimian taxa that are also viable contenders for strepsirhine status. Plesiopithecus teras is a Nycticebus-sized, nocturnal prosimian from the late Eocene, Fayum, Egypt, that shares cranial specializations with lorisoids, but it also retains primitive features (e.g. four premolars) and has unique specializations of the anterior teeth excluding it from direct lorisiform ancestry. Another unnamed Fayum primate resembles modern cheirogaleids in dental structure and body size. Two genera from Oman, Omanodon and Shizarodon, also reveal a mix of similarities to both cheirogaleids and anchomomyin adapoids. Resolving the phylogenetic position of these Africa primates of the early Tertiary will surely require more and better fossils. By the early to middle Miocene, lorisoids were well established in East Africa, and the debate about whether these represent lorisines or galagines is reviewed. Neontological data are used to address the controversial branching sequences among extant lorisid clades. Data from the skin and scent glands, when integrated with other lines of evidence, suggest that Asian and African lorisines share a common lorisine ancestry. The hypothesis of an African clade containing both pottos and galagos to the exclusion of Asian lorisines is less tenable. True galagines are found in the fossil record of Namibia, while true lorisines are known from the Miocene of Asia. The hypothetical branching sequences can be integrated with behavioral and morphological features to develop an adaptive model of lorisoid divergence. By specializing on two different foraging modes early in their radiation, lorisines and galagines subsequently underwent a chain of integrated evolutionary changes eventually having an impact on many components of locomotor behavior, anatomy, physiology, reproduction, life history, and social behavior. Ongoing evolutionary studies of extant galagines are illuminating population phenomena and processes of speciation in an ecological context.

  Website 

Book chapters

• Bearder S, Nekaris K, Lloyd J, 'Tips from the bush: an A-Z of suggestions for successful fieldwork' in Field and Laboratory Methods in Primatology: A Practical Guide, Cambridge University Press (2011)
  ISBN: 9780521142137
  Abstract Building on the success of the first edition and bringing together contributions from a range of experts in the field, the second edition of this guide to research on wild primates covers the latest advances in the field, including new information on field experiments and measuring behaviour. It provides essential information and advice on the technical and practical aspects of both field and laboratory methods, covering topics such as ethnoprimatology; remote sensing; GPS and radio-tracking; trapping and handling; dietary ecology; and non-invasive genetics and endocrinology. This integrated approach opens up new opportunities to study the behavioural ecology of some of the most endangered primates and to collect information on previously studied populations. Chapters include methodological techniques; instructions on collecting, processing and preserving samples/data for later analysis; ethical considerations; comparative costs; and further reading, making this an invaluable tool for postgraduate students and researchers in primatology, behavioural ecology and zoology.
• Nekaris K, Starr C, Collins R, Wilson A, 'Comparative Ecology of Exudate Feeding by Lorises (Nycticebus, Loris) and Pottos (Perodicticus, Arctocebus)' in The Evolution of Exudativory in Primates, Springer (2010)
  ISBN: 9781441966605
  Abstract

  The Evolution of Exudativory in Primates is the first edited volume to offer a comprehensive overview of this rare dietary niche in the primate order. Leading researchers in the field of primatology synthesize our current knowledge of the behavioral, socioecological, nutritional, morphological, and evolutionary aspects of exudate-feeding in primates. The Evolution of Exudativory in Primates covers exudate-feeding in callitrichines, callimicos, mouse lemurs, lorises, and galagos. Advances in our understanding of how these animals obtain their food and digest it, how this food resource affects social relationships, and how morphology is related to exudate-feeding are presented in subsequent essays. The final chapter synthesizes current data on what role exudate-feeding may have played in the earliest primates, the plesiadapiforms, and what exudate-feeding signals may be present in the fossil record. Ideal for upper-level undergraduate and graduate primatology courses, The Evolution of Exudativory in Primates can also be used for courses in biology, comparative mammalogy, and conservation. About the Editors: ANNE M. BURROWS is an Associate Professor in the Department of Physical Therapy at Duquesne University and a Research Associate Professor in the Department of Anthropology at the University of Pittsburgh. She has worked on the functional and evolutionary morphology of the primate craniofacial complex with a focus on strepsirrhines. LEANNE T. NASH is a Professor of Anthropology in the School of Human Evolution and Social Change at Arizona State University. She has worked extensively with a captive colony of Galago senegalensis held previously at ASU for 20 years. She has also done fieldwork in Africa and Madagascar on baboons, galagos, and sportive lemurs. Other collaborations have been on captive chimpanzee behavior with the Primate Foundation of Arizona.

  Website 
• Nekaris K A I, Bearder S K, 'The Lorisiforme primates of Asia and mainland Africa: diversity shrouded in darkness' in Primates in Perspective, Oxford University Press (2010)
  ISBN: 9780195390438
  Abstract Featuring forty-seven original essays by seventy leading researchers, Primates in Perspective, Second Edition, offers a comprehensive and contemporary overview of all major areas of primatology. Thoroughly revised and updated throughout, the second edition offers a diversity of theoretical positions on such topics as reproduction, ecology, and social behavior and intelligence. Primates in Perspective, Second Edition, is ideal for introductory primatology courses and can also be used in upper-division behavior and conservation courses. Additionally, it is an essential reference for primate researchers.
• Nekaris K, Munds R, 'Using Facial Markings to Unmask Diversity: The Slow Lorises (Primates: Lorisidae: Nycticebus spp.) of Indonesia' in , Springer (2010)
  Abstract The slow lorises (Nycticebus) are the only strepsirrhine primates found in Indonesia (Nekaris and Bearder 2007). In addition to features such as a toothcomb and moist nose, these small nocturnal primates were given their name based on their trademark steady, stealthy, and fluid locomotion. Morphologically incapable of leaping (Sellers 1996), slow lorises rather slither through the treetops, and if startled, they may freeze or even cover their face, resulting in one of their many Indonesian names, malu malu or -œthe shy one- (Supriatna and Wahyono 2000). Alternatively, they can fleetingly but silently escape, resulting in the name buah angin or -œwind monkey- in Acehnese (Nekaris and Nijman 2007a). One of two genera of nocturnal primates found in Indonesia (the other being Tarsius), slow lorises are a unique part of Indonesian primate communities, and are widely spread on at least 27 of Indonesia" s islands, including Borneo, Sumatra, and Java (Table 22.1) (Nijman and Nekaris in press). Despite this, studies of Indonesian slow lorises are in their infancy. Website 
• Nekaris K, Pimley E, Ablard K, 'Predator Defense by Slender Lorises and Pottos' in Primate Anti-Predator Strategies, Springer (2007)
  ISBN: 9781441941909
  Abstract Since the 1960s, primatologists have recognized the impact of predation on the evolution of morphology, the social systems and cognitive behavior of monkeys and apes, but few studies considered its impact on the prosimians - lemurs, lorises, galagos and tarsiers. This comprehensive volume, written by experts in the field, narrows this gap by highlighting the effect of predation on the order Primates in general. Theoretical approaches to understanding how primates perceive predation threat, as well as proximate and ultimate causes to address threat and attack, are considered across the primate order. Although this volume concentrates on the least known group in this theoretical area - the prosimians - contributions by researchers on numerous primate taxa across four major geographical regions make this a novel and exciting contribution to students interested in primate evolution and ecology.
• Nekaris K, Gursky S, 'Theoretical perspectives of primate anti-predator behaviour' in Primate Anti-Predator Strategies, Springer (2007)
  ISBN: 9781441941909
  Abstract Since the 1960s, primatologists have recognized the impact of predation on the evolution of morphology, the social systems and cognitive behavior of monkeys and apes, but few studies considered its impact on the prosimians - lemurs, lorises, galagos and tarsiers. This comprehensive volume, written by experts in the field, narrows this gap by highlighting the effect of predation on the order Primates in general. Theoretical approaches to understanding how primates perceive predation threat, as well as proximate and ultimate causes to address threat and attack, are considered across the primate order. Although this volume concentrates on the least known group in this theoretical area - the prosimians - contributions by researchers on numerous primate taxa across four major geographical regions make this a novel and exciting contribution to students interested in primate evolution and ecology.
• Bearder S, Nekaris K, Buzzell C, 'Dangers in the Dark: are some noctural primates afraid of the dark?' in Eat or be Eaten: Predator Sensitive Foraging Among Primates, Cambridge University Press (2002)
  ISBN: 9780521011044
  Abstract This volume brings together primary data from a variety of primate species living in both natural habitats and experimental settings, and explores the variables that may play a role in primates' behavioral strategies. Taken together, these studies demonstrate that predator sensitive foraging is relevant to many primates, of various body sizes and group sizes and living in different environments. Eat or be Eaten encourages further discussion and investigation of the subject and will make fascinating reading for researchers and students in primatology, ecology, and animal behavior.Website 

Other publications

• Lehtinen J, Nekaris KAL, Nijman V, Coudrat C, Wirdateti, 'Distribution of the Javan Slow Loris (Nycticebus Javanicus): Assessing the Presence in East Java, Indonesia'
  Folia Primatologica 84 (2013) pp.295-295ISSN: 0015-5713 eISSN: 1421-9980
  
• Nijman V, Nekaris KAI, 'Assessing Conflict Between Humans and Commensal Non-human Primates in Sri Lanka Following An Ethnoprimatological Approach'
  Folia Primatologica 84 (2013) pp.307-308ISSN: 0015-5713 eISSN: 1421-9980
  
• Nekaris KAI, Pambudi JAA, Ahmad RD, Susanto D, Raharjo B, Nijman V, 'Densities, Distribution and Detectability of a Small Nocturnal Primate (javan Slow Loris Nycticebus Javanicus) in a Montane Rainforest'
  Folia Primatologica 84 (2013) pp.306-306ISSN: 0015-5713 eISSN: 1421-9980
  
• Munds RA, Nekaris KAI, Nijman V, Goossens B, 'Living Together in the Night: Abundance and Habitat Use of Sympatric and Allopatric Populations of Slow Lorises and Tarsiers (Nycticebus and Tarsius).'
  American Journal of Physical Anthropology 150 (2013) pp.203-203Abstract

  Throughout much of Asia slow lorises (Nycticebus) and tarsiers (Tarsius) live allopatrically but on several islands they occur in sympatry. As habitats dwindle, competition for resources may increase within the area of sympatry. An understanding of how they are coping with competition is necessary for conservation measures. To address this we gathered data on the abundances and vertical strata preferences of slow loris and tarsier species from the literature. We predict sympatric species will favour different heights from congeners. Allopatric species will have lower abundances compared to sympatric species. In addition, we studied tarsiers and lorises in sympatry in Sabah, Borneo. We estimated abundances of Bornean lorises (Nycticebus menagensis) and Western tarsiers (Tarsius bancanus borneanus) and investigated habitat use. Through the literature review we found lorises do not vary in densities, whether allopatric or sympatric. Abundances of sympatric and allopatric tarsiers were significant (sympatric: 3-27 individuals/km², allopatric: 57-268 individuals/ km²). Vertical strata use of sympatric and allopatric tarsier populations was not significant, but was for sympatric and allopatric loris populations (p=0.036). On a small scale estimated densities of Bornean lorsies were 5.105 slow lorises/km² and 3.646 individuals/km² for Western tarsiers. Slow lorises favoured the upper and middle level of the forest (10-30 m) and tarsiers the lower levels (<5 m). In our analysis of vegetation plots we found that tree heights and diameter at breast height preferences differed between genera. Results indicate sympatric genera are able to share their nocturnal environment due to niche separation.

  Website 
• Nijman V, Pambudi JA, Achmed D, Nekaris KA, 'Densities, Distribution and Detectability of a Small Nocturnal Primate (Javan Slow Loris Nycticebus Javanicus) in a Montane Rainforest'
  American Journal of Physical Anthropology 150 (2013) pp.207-207Abstract
  Nocturnal mammals can be challenging to survey and, especially for many species
  that live in dense forest habitats, limited information is available on densities and distributions. We surveyed the endemic Javan slow loris Nycticebus javanicus in the montane forests of Mount Gede Pangrango, West Java, Indonesia. Surveys were conducted on 23 transects (260 h covering some 93 km) walking at variable speeds between 200 and 800 m h−1. Densities on individual transects varied from 0 to 52 ind. km−2, with an overall density of 15.6 ind. km−2 (95% CI 9.7 to 25.2 ind. km−2). Encounter rates per kilometre were strongly influenced by the speed at which transects were walked, with fewer lorises detected at higher speeds. This effect was absent when considering encounter rates per hour. Detectability and behavior of Javan slow lorises were not affected by the amount of lunar light and, in contrast to studies of some of their congeners, we found no evidence of lunar phobia or lunar philia. Our study shows that slow lorises are not homogeneously distributed in their montane habitat and occur at intermediate densities. However, encounter rates did not differ between disturbed and primary forest. Analysis of data from multiple surveys of lorisforms, including this one, reveals no statistically significant relationship between survey effort or the speed at which transects are walked and estimated densities, but speed is positively correlated with encounter rates.
• Nijman V, Nekaris KAI, 'Diurnal Eagles As Predators of Primates in Asia'
  Folia Primatologica 82 (2012) pp.355-356ISSN: 0015-5713 eISSN: 1421-9980
  
• Nijman V, Nekaris KAI, Bickford DP, 'Asian Medicine: Small Species at Risk'
  Nature 481 (2012) pp.265-265ISSN: 0028-0836
  
• Svensson MS, Nijman V, Nekaris KAI, 'In Situ and Ex Situ Conservation of African Pottos and Angwantibos: Where Are We and Where Do We Need to Go?'
  Folia Primatologica 82 (2012) pp.390-390ISSN: 0015-5713 eISSN: 1421-9980
  Abstract

  Despite a large geographic distribution, the African nocturnal perodicticines, pottos (Perodicticus) and angwantibos (Arctocebus ) remain amongst the least studied primate taxa. Only two one-year field studies have been conducted on pottos, with only circumstantial data onangwantibos, coming from field data on shot animals. Through a meta-analysis, based on published literature and unpublished reports, we here review recent skeletal and genetic evidence that point to five species, and indicate far more diversity: Perodicticus potto, P. Ibeanus, P. edwardsi, Arctocebus aureus, A. calabarensis. Pelage colour, dorsal markings and tail characteristics (for pottos) supplement the genetic data that suggest these comprise distinct taxa. Arctocebus ranges from Nigeria and Cameroon in the north to Gabon and Congo in the south, whilst Perodicticus is more widespread, having a combined distribution that encompasses a large portion of central and western Africa. Despite this large range, only nine surveys with any substantial effort have been published in the last twenty years from only five range countries; two re-ported new taxa (P. p. stockleyi and P. p. juju). Although not included in abundance counts,pottos frequently appear in bushmeat reports throughout their range, suggesting this is a realthreat to this cryptic, easy-to-catch primate. Despite an almost complete dearth of knowledgeabout these taxa, all perodicticine taxa are listed on the IUCN Red List of Threatened Species as Least Concern, with, contradictorily, only the reasonably studied P. p. stockleyi as Data Deficient. The situation of these primates is mirrored in zoos. No Arctocebus are kept in European zoos, with only 17 Perodicticus in ISIS institutions. Their taxonomy is uncertain, breeding rates are poor and infant mortality is high. Clearly the perodictines offer an open frontier for both in situ and ex situ studies. We certainly do not have the data to assess their conservation status yet.

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Further information

Related Links

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Education

• PhD and MA in Biological Anthropology, Washington University, St Louis.
• Certificat de Primatologie, Universite Louis Pasteur, Strasbourg, France.
• BA in Biological Anthropology, University of Missouri, Columbia, USA.

Current positions

• Professor in Primate Conservation
• Course Tutor MSc Primate Conservation