Sensitivity of Vanessa cardui to Temperature Variations: A Cost-Effective Experiment for Environmental Education.

Temperature increases mediated through climate change threaten the survival of species. It is of foremost importance to engage citizens and future generations in understanding the mechanisms through which temperatures impose their effects. For educators, this is not straightforward, as tools for examining the impact of temperature over the lifetime of an animal are prohibitively expensive. At the same time, environmental educators need guidance on the appropriate study systems to use with a balance between the species having an obvious response and ensuring the outcomes are ethical and sustainable. In our study, we created and tested a cost-effective experiment meant to be used for environmental education purposes. More specifically, we tested the sensitivity of the painted lady butterfly Vanessa cardui to temperature variations using a homemade incubator. We describe the design of this experiment and report findings on survival rate, morphological variations, development time of various stages and wingspan of adults across a range of biologically relevant temperatures. The information provided gives educators options for testing a variety of hypotheses with regards to the impacts of temperature using an affordable and flexible set-up. Furthermore, the findings can be used by students to develop an understanding of the ramifications of the butterflies' responses in an ecological context.

Effect of shade on biodiversity within coffee farms: A meta-analysis.

Aligning crop production with conservation initiatives has long been a topic of debate, with agricultural intensification threatening biodiversity across the globe. Shade-grown coffee allows farmers to preserve biodiversity by providing viable habitat, but its conservation value remains unclear. In this meta-analysis, we screened existing literature using the PRISMA protocol to compare the effect of three shade intensities on species diversity and individual abundance: sun, low shade (LS) and high shade (HS). Furthermore, we examine differences between taxa, within taxa and between regions to establish which species benefit most from shade and whether these benefits vary dependent on geographical location. Out of 1889 studies, we included 69 studies in the analysis, and performed random-effects meta-analyses and meta-regressions. Overall, we found that species diversity was significantly higher in HS when compared to sun and LS, and species diversity in LS tended to be higher than in sun. In each treatment, the species diversity of birds was higher in the higher shade treatment, i.e., HS and LS. In addition, mammal and epiphyte species diversity was higher in HS when compared to LS. Similarly, studies from Latin America showed significantly higher species diversity and abundance in shaded farms when compared to sun farms. Studies conducted in Africa detailed the opposite relationship, with abundance being significantly higher in less shaded systems, highlighting that land-use strategies must be region-specific. Moving forward, strategies to conserve biodiversity within coffee farms should: 1) account for region-specific variables; 2) end further encroachment; 3) maintain connectivity; and 4) optimise yield through prioritising faunal and floral diversity.

COVID-19's Impact on the Pan African Sanctuary Alliance: Challenging Times and Resilience from Its Members.

The worldwide pandemic caused by SARS-CoV-2 challenged conservation organizations. The lack of tourism has benefited or negatively affected wildlife organizations in various ways, with several primate sanctuaries struggling to cope with the COVID-19 crisis and to keep providing for their inhabitants. In addition, the genetic similarity between great apes and humans puts them at higher risk than any other species for the transmission of COVID-19. PASA is a non-profit organization comprising 23 sanctuaries, and cares for many species of primate, including African great apes. In light of the pandemic, we aimed to understand the direct effects of COVID-19 on PASA management throughout three time periods: before (2018-2019), at the start of (2019-2020), and during (2020-2021) the pandemic. We collected data via annual surveys for PASA members and ran Generalized Linear Mixed Models to highlight any significant differences in their management that could be linked to COVID-19. Our findings demonstrated no particular impact on the number of primates rescued, employees, or expenses. However, revenues have been decreasing post-COVID-19 due to the lack of income from tourism and volunteer programs. Nonetheless, our results reveal a form of resilience regarding the sanctuaries and the strategy applied to maintain their management. Consequently, we emphasize the specific impacts of the COVID-19 outbreak and its repercussions for conservation work. We discuss the difficulties that sanctuaries have faced throughout the crisis and present the best measures to prevent future outbreaks and protect biodiversity.

Road Infrastructure and Primate Conservation: Introducing the Global Primate Roadkill Database.

As road infrastructure networks rapidly expand globally, especially in the tropics, previously continuous habitats are being fragmented, resulting in more frequent wildlife-vehicle collisions (WVC). Primates are widespread throughout many sub-/tropical countries, and as their habitats are fragmented, they are increasingly at risk of WVC. We created the Global Primate Roadkill Database (GPRD), the largest available standardized database of primate roadkill incidents. We obtained data from published papers, un-published and citizen science databases, anecdotal reports, news reports, and social media posts. Here, we describe the collection methods for the GPRD and present the most up-to-date version of the database in full. For each primate roadkill incident, we recorded the species killed, the exact location, and the year and month the roadkill was observed. At the time of publication, the GPRD includes 2862 individual primate roadkill records from 41 countries. As primates range in more than twice as many countries, the absence of data from these countries is not necessarily indicative of a lack of primate vehicular collisions. Given the value of these data for addressing both local and global research questions, we encourage conservationists and citizen scientists to contribute to the GPRD so that, together, we can better understand the impact road infrastructure has on primates and evaluate measures which may help mitigate risk-prone areas or species.

Applications of Accelerometers and Other Bio-Logging Devices in Captive and Wild Animals.

Bio-logging devices have been widely used in ecology across a range of species to acquire information on the secret lives of animals in the wild, which would otherwise be challenging to obtain via direct observations [...].

It's a sign: Animal welfare and zoo type are predictors of animal identification signage usage and quality at zoo exhibits.

Conservation education programs are listed as priority actions for almost every threatened species on the International Union for Conservation of Nature Red List. Zoos play an important role in delivering such programs, yet evidence of zoo education in many non-western countries is limited. Here, we evaluate animal identification signage prevalence and quality at zoo exhibits and investigate whether animal welfare, zoo type (accredited, government, and private), admission fee, zoo size, and proximity to urban centers are influencing factors. We used hornbills (Bucerotidae) as a model taxon, surveyed hornbill signage, and conducted welfare assessments of hornbill exhibits. We developed scoring frameworks and applied content analysis to analyze signage quality. Our results show that out of 18 zoos that displayed hornbills, 15 had hornbill signage. However, of the 106 hornbill exhibits in these zoos, 33% had no signage. We also found that signage presence or absence at individual zoos and signage quality is strongly correlated with animal welfare quality. Zoo type is a key factor in predicting signage and welfare quality, with accredited zoos scoring highest for both signage and welfare, followed by government and private zoos. Private zoos charged higher admission than other zoo types, and zoo size and proximity to urban centers did not influence signage or welfare scores. Overall, we conclude that in our study, signage usage and quality are inadequate, highlighting the importance of compliance with robust zoo standards to improve education and welfare within zoos to support global conservation goals.

Risk of Viral Infectious Diseases from Live Bats, Primates, Rodents and Carnivores for Sale in Indonesian Wildlife Markets.

Southeast Asia is considered a global hotspot of emerging zoonotic diseases. There, wildlife is commonly traded under poor sanitary conditions in open markets; these markets have been considered 'the perfect storm' for zoonotic disease transmission. We assessed the potential of wildlife trade in spreading viral diseases by quantifying the number of wild animals of four mammalian orders (Rodentia, Chiroptera, Carnivora and Primates) on sale in 14 Indonesian wildlife markets and identifying zoonotic viruses potentially hosted by these animals. We constructed a network analysis to visualize the animals that are traded alongside each other that may carry similar viruses. We recorded 6725 wild animals of at least 15 species on sale. Cities and markets with larger human population and number of stalls, respectively, offered more individuals for sale. Eight out of 15 animal taxa recorded are hosts of 17 zoonotic virus species, nine of which can infect more than one species as a host. The network analysis showed that long-tailed macaque has the greatest potential for spreading viral diseases, since it is simultaneously the most traded species, sold in 13/14 markets, and a potential host for nine viruses. It is traded alongside pig-tailed macaques in three markets, with which it shares six viruses in common (Cowpox, Dengue, Hepatitis E, Herpes B, Simian foamy, and Simian retrovirus type D). Short-nosed fruit bats and large flying foxes are potential hosts of Nipah virus and are also sold in large quantities in 10/14 markets. This study highlights the need for better surveillance and sanitary conditions to avoid the negative health impacts of unregulated wildlife markets.

Factors influencing terrestriality in primates of the Americas and Madagascar.

Among mammals, the order Primates is exceptional in having a high taxonomic richness in which the taxa are arboreal, semiterrestrial, or terrestrial. Although habitual terrestriality is pervasive among the apes and African and Asian monkeys (catarrhines), it is largely absent among monkeys of the Americas (platyrrhines), as well as galagos, lemurs, and lorises (strepsirrhines), which are mostly arboreal. Numerous ecological drivers and species-specific factors are suggested to set the conditions for an evolutionary shift from arboreality to terrestriality, and current environmental conditions may provide analogous scenarios to those transitional periods. Therefore, we investigated predominantly arboreal, diurnal primate genera from the Americas and Madagascar that lack fully terrestrial taxa, to determine whether ecological drivers (habitat canopy cover, predation risk, maximum temperature, precipitation, primate species richness, human population density, and distance to roads) or species-specific traits (body mass, group size, and degree of frugivory) associate with increased terrestriality. We collated 150,961 observation hours across 2,227 months from 47 species at 20 sites in Madagascar and 48 sites in the Americas. Multiple factors were associated with ground use in these otherwise arboreal species, including increased temperature, a decrease in canopy cover, a dietary shift away from frugivory, and larger group size. These factors mostly explain intraspecific differences in terrestriality. As humanity modifies habitats and causes climate change, our results suggest that species already inhabiting hot, sparsely canopied sites, and exhibiting more generalized diets, are more likely to shift toward greater ground use.

Physiological Indicators of Acute and Chronic Stress in Securely and Insecurely Attached Dogs Undergoing a Strange Situation Procedure (SSP): Preliminary Results.

The quality of the attachment bond towards the caregiver may affect the dog's physiological responses to stressful stimuli. This study aimed to measure chronic and acute physiological parameters of stress in ten securely and ten insecurely attached dogs. The twenty experimental subjects were selected from a sample of dogs that participated with their owners in the Strange Situation Procedure. Saliva samples were collected before (T0) and after (T1) the test. Blood pressure, heart rate, respiratory rate, and rectal temperature were measured after the test, only. At this time, a hair sample was also collected. RM ANOVA was used to analyse cortisol concentrations between secure and insecure dogs at T0 and T1. Mann-Whitney U test or T test were used for other physiological parameters. Insecure dogs had significant higher salivary cortisol concentrations than secure dogs at T1 (p = 0.024), but only a non-significant trend towards higher cortisol concentrations at T0 (p = 0.099). Post-test heart rate also tended to be higher in insecure compared to secure dogs (p = 0.077). No significant differences in hair cortisol concentration were found. The quality of attachment may affect the dog's physiological response to acute stress, at least when related to separation from the caregiver. The effect of attachment on chronic stress requires further investigation.

The Use of Live Action, Animation, and Computer-Generated Imagery in the Depiction of Non-Human Primates in Film.

For over 100 years, non-human primates (primates) have been a part of the now hundred-billion-dollar global film industry in a variety of capacities. Their use in the film industry is of concern due to the negative welfare effects on individuals, the potential for increased pet trade, and the conservation impacts of public perception. While the effects on human perception of using live primates in film have been studied, little research has been performed on their appearance in animation and none in computer-generated imagery (CGI). We aimed to investigate how the portrayal of primates varied between depiction medium types and how this related to the films' performance with critics and in the box office. We observed 151 primates in 101 different English-speaking films that debuted between 2000 and 2019. For each appearance we recorded aspects of primate portrayals based on accuracy, anthropomorphism, environment, and agency displayed, along with the depiction medium. We used structural equation models to depict the highest likelihood of the portrayal aspects on the medium's relationship to the films gross profit worldwide and film critic consensus scores. We found that over the 20-year time frame, use of live primates has decreased, CGI has increased, and animations have remained relatively steady. While animation had no significant relationship to gross profit or critic consensus, both were significantly lower for films that used live primates and were significantly higher for films that used CGI primates. Due to the steady increase in the use of the CGI medium and its positive relationship with gross profit and critic consensus, it could have great effects on people's perceptions of primates and implications for conservation efforts.

Disentangling the Legal and Illegal Wildlife Trade-Insights from Indonesian Wildlife Market Surveys.

It is challenging to disentangle the legal and illegal aspects of wild-caught animals that are traded in wildlife markets or online, and this may diminish the value of conducting wildlife trade surveys. We present empirical studies on the trade in birds (ducks, owls, songbirds, non-passerines) in Indonesia (2005 to 2021). Based on visits to wildlife markets, wholesale traders, and monitoring of an Instagram account, we examine if five specific pieces of legislation (domestic and international) are adhered to: (1) protected species, (2) harvest quota, (3) welfare, (4) provincial transport restrictions, and (5) illegal import of CITES-listed species. Our five distinctly different case studies showed that in each case, certain rules and regulations were adhered to, whilst others were violated to varying degrees. When trade involved non-protected species, there was frequently a lack of harvest quotas or trade occurred above these allocated quotas. Basic welfare provisions were regularly and habitually violated. Visiting wildlife markets and recording first-hand what is openly offered for sale is a highly reliable, verifiable, and valuable method of data collection that can give insight in numerous aspects of the animal trade. Our research provides support for recognising the urgency for the government to take appropriate action to curb all the illegal aspects of the bird trade in Indonesia.

Training in the Dark: Using Target Training for Non-Invasive Application and Validation of Accelerometer Devices for an Endangered Primate (Nycticebus bengalensis).

Accelerometers offer unique opportunities to study the behaviour of cryptic animals but require validation to show their accuracy in identifying behaviours. This validation is often undertaken in captivity before use in the wild. While zoos provide important opportunities for trial field techniques, they must consider the welfare and health of the individuals in their care and researchers must opt for the least invasive techniques. We used positive reinforcement training to attach and detach a collar with an accelerometer to an individual Bengal slow loris (Nycticebus bengalensis) at the Shaldon Wildlife Trust, U.K. This allowed us to collect accelerometer data at different periods between January-June 2020 and January-February 2021, totalling 42 h of data with corresponding video for validation. Of these data, we selected 54 min where ten behaviours were present and ran a random forest model. We needed 39 15-min sessions to train the animal to wear/remove the collar. The accelerometer data had an accuracy of 80.7 ± SD 9.9% in predicting the behaviours, with 99.8% accuracy in predicting resting, and a lower accuracy (but still >75% for all of them apart from suspensory walk) for the different types of locomotion and feeding behaviours. This training and validation technique can be used in similar species and shows the importance of working with zoos for in situ conservation (e.g., validation of field techniques).

Bird Assemblages in Coffee Agroforestry Systems and Other Human Modified Habitats in Indonesia.

Deforestation in the tropics is mainly driven by the need to expand agriculture and forestry land. Tropical cropland has also undergone a process of intensification, particularly evident in regions that are the main exporters of deforestation-driven commodities. Around 25 million people in the world depend on coffee production, which has a profound contribution to global biodiversity loss through agricultural extensification and intensification. Nevertheless, coffee agroforestry systems have been postulated to serve as an alternative refuge for biodiversity across different regions. We aim to compare bird abundance, diversity, and richness in commercial polyculture coffee systems (i.e., the highest degree of habitat complexity that can be achieved in coffee fields after deforestation) with other coffee agroforestry systems and human modified habitats in Java, Indonesia. We collected data in 21 sites (1228 points) on Java from February to August 2021 using the point sampling method. Via generalised additive models, we tested whether the abundance, diversity, and richness of birds were different between different human modified habitats including other potential predictors such as elevation, distance to protected areas, shade tree richness, and plant diversity. Using the non-metric multidimensional scaling, we tested whether there was a difference in terms of the composition of foraging guilds between habitats. Commercial polyculture coffee fields can sustain levels of bird abundance, diversity, and richness comparable to agroforestry systems under natural forest, and higher than sun coffee and shaded monoculture coffee, and of other human modified habitats such as crop/fruit fields and tree farms. Coffee agroforestry systems have a higher proportion of nectarivores, insectivores, and frugivores than other systems that can sustain high diversity and richness of birds such as paddy fields that mainly have granivores and carnivores. Complex polycultures can represent an avenue for the future of sustainable agriculture in conditions where deforestation rates are high and in crops such as coffee, which maintain high yield in the presence of diverse shade.

The sticky tasty: the nutritional content of the exudativorous diet of the Javan slow loris in a lowland forest.

Plant exudates are an important food source for many primates. The Critically Endangered Javan slow loris (Nycticebus javanicus) was previously found to prefer Acacia decurrens exudate in an anthropogenically disturbed site, while its feeding habits in secondary natural forest remain unknown. Knowledge of the chemical characteristics of the plant exudates that Javan slow lorises consume is limited, especially with respect to those that they feed on in natural forests. As plant exudates may contain plant secondary metabolites (PSM), which are considered unpalatable in high concentrations, differences in PSM composition may drive feeding preferences. This research aims firstly to confirm exudate consumption by the Javan slow loris in a lowland tropical forest in Central Java, and secondly to identify the chemical characteristics of the exudates consumed. We followed wild slow lorises in Kemuning Forest, Central Java and observed their behaviour. We investigated the gum-producing trees that were utilized by the slow lorises by tapping the exudates and examining their nutritional and PSM contents. We found that exudates are the predominant food source for the Javan slow loris in this lowland forest, and that their nutritional contents are similar to those of exudates consumed by lorises in anthropogenically disturbed areas. Significant differences in polysaccharide and flavonoid contents were found between consumed and unconsumed exudates. Knowledge of the diet of the Javan slow loris is crucial to its conservation, and our findings confirm the importance of exudates in its diet. We also highlight the need to preserve natural slow loris habitat, and to manage the diets of these species in captivity. The results of this study indicate that plant exudates should constitute a significant portion of the diet of captive slow lorises, and that the presence of exudate-producing trees is vital in areas into which slow lorises are to be translocated.

Silky Sifakas (Propithecus candidus) Use Sleep Sites for Thermoregulation, Food Access and Predator Avoidance.

Primate sleeping site selection is influenced by multiple ecological factors including predation avoidance, thermoregulation and food access. To test these hypotheses, we studied the sleeping trees used by a group of wild silky sifakas (Propithecus candidus) in Marojejy National Park, Madagascar. During this 10-month study, the group slept in 828 sleeping trees from approximately 35 genera. In support of thermoregulation, generalized linear models revealed that as temperature decreased, the number of individuals sleeping together significantly increased and they slept at further distances from the trunk. As rainfall increased, sleep site height significantly increased. Weinmannia was the most frequented tree genus, despite low abundance, accounting for 29% of all sleeping trees. In support of food access, 94.8% of sleeping trees were food trees. Weinmannia is among the most highly preferred food trees. The group slept at a mean height of 16.0 m near the top of tall trees which averaged 19.5 m. Sleep trees were significantly taller than trees in botanical plots within the sifaka's home range. They never slept in the same trees on consecutive nights, and sleeping heights were significantly higher than daytime heights which is consistent with predation avoidance. Social sleeping in groups of 2 or 3 individuals (62.9%) was more common than solitary sleeping (37.1%). At such heights, huddling may increase vigilance and lessen the risk of predation by fossa (Cryptoprocta ferox) while also reducing heat loss. These patterns suggest that silky sifaka sleep site choice is influenced by thermoregulation and food access in addition to predation avoidance. We suggest that understanding sleep site use can assist in conservation of species like silky sifakas by enabling researchers to find new groups, protect habitats with key tree species and inform reforestation efforts.

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

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

The Role of the Songbird Trade as an Anthropogenic Vector in the Spread of Invasive Non-Native Mynas in Indonesia.

The wildlife trade has facilitated the introduction of invasive non-native species, which may compete with native species for resources and alter ecosystems. Some of these species have great potential to become invasive if released or escaped from captivity. Here we studied the pet trade in a group of open countryside birds, the mynas (Acridotheres spp.) in Indonesia, and identified the areas that are at high risk of facing the establishment of these species. Mynas are among the most invasive birds in Southeast Asia. Once established in a new area, they are almost impossible to eradicate and can have strong negative impacts on the ecosystem. Preventing their introduction is therefore essential. Yet, invasive non-native mynas continue to be traded openly. We present data on the trade in seven species of mynas on Java, Bali and Lombok, with three species being native to parts of one or two of these islands, but not to the remainder, and four that are non-native to the region. From 2016 to 2021 we conducted 255 surveys of 30 animal markets. We recorded over 6000 mynas that were offered for sale outside their native range. Areas most at risk because of their high prevalence in specific animal markets, are Greater Jakarta, eastern Java, Bali and Lombok. The number of invasive non-native mynas recorded was positively related to the size of the animal market. Indonesia is signatory to several international agreements (CBD, ASEAN) that have policies and guidelines to prevent the introduction of invasive non-native species, but compliancy is weak. Annually hundreds and possibly thousands of invasive non-native mynas are released by Indonesian conservation authorities in regions that are outside their native range. Effective management of, and regulation of trade in, potential invasive non-native birds in Indonesia falls short and inadvertently greatly aids both their introduction and establishment.

The Influence of Seasonal Availability of Young Leaves on Dietary Niche Separation in Two Ecologically Similar Folivorous Lemurs.

Traditional socio-ecological models consider that folivorous primates experience limited feeding competition due to the low quality, high abundance, and even distribution of leaves. Evidence from several folivorous species that experience similar constraints to frugivores does not support this hypothesis. The sympatric lemur genera Avahi (Indriidae) and Lepilemur (Lepilemuridae) are good models to understand how food availability constrains folivores since they are both nocturnal, folivorous, and have a comparable body mass. Here we investigate how two nocturnal folivorous primates, Avahi meridionalis and Lepilemur fleuretae, living in the lowland rain forest of Tsitongambarika, South-East Madagascar, partition their dietary niche and are influenced by seasonality of young leaves. To account for food availability, we collected annual phenological data on 769 trees from 200 species. We also collected behavioural data on 5 individuals per lemur species from August 2015 to July 2016 via continuous focal sampling. We found the phenological profile to be seasonal with peaks of leaf flushing, flowering, and fruiting occurring in the austral summer. The two species showed limited dietary overlap (37% rich period, 6% lean period), and A. meridionalis showed higher feeding time and longer daily distances travelled during the rich period. Lepilemur fleuretae showed a dietary shift during the lean period, relying more on mature leaves (73.3% during the lean period, 13.5% during the rich period) but maintaining similar activity levels between seasons. The time spent feeding on food items by A. meridionalis was positively correlated with the nitrogen content and negatively correlated with polyphenols during the rich period. We highlighted a clear effect of the seasonality of young leaves on the diet, nutritional content, activity patterns, and daily distances travelled by two folivorous species, which can be linked to nutrient balancing and time-minimising versus energy-maximising strategies.

Risky Business: The Function of Play in a Venomous Mammal-The Javan Slow Loris (Nycticebus javanicus).

Immature mammals require opportunities to develop skills that will affect their competitive abilities and reproductive success as adults. One way these benefits may be achieved is through play behavior. While skills in developing use of tusks, antlers, and other weapons mammals have been linked to play, play in venomous animals has rarely been studied. Javan slow lorises (Nycticebus javanicus) use venom to aid in intraspecific competition, yet whether individuals use any behavioral mechanisms to develop the ability to use venom remains unclear. From April 2012 to December 2020, we recorded 663 play events and studied the factors influencing the frequency of play and the postures used during play in wild Javan slow lorises. Regardless of the presence of siblings, two thirds of play partners of young slow lorises were older and more experienced adults. Young lorises engaged in riskier behaviors during play, including using more strenuous postures and playing more in riskier conditions with increased rain and moonlight. We found that play patterns in immature lorises bear resemblance to venom postures used by adults. We suggest that play functions to train immature lorises to deal with future unexpected events, such as random attacks, as seen in other mammalian taxa with weapons. Given the importance of venom use for highly territorial slow lorises throughout their adult lives and the similarities between venom and play postures, we cannot rule out the possibility that play also prepares animals for future venomous fights. We provide here a baseline for the further exploration of the development of this unique behavior in one of the few venomous mammals.

Slow lorises use venom as a weapon in intraspecific competition.

Animals have evolved an array of spectacular weapons, including antlers, forceps, proboscises, stingers, tusks and horns [1]. Weapons can be present in males and females of species needing to defend critical limiting resources, including food (rhinoceros beetles, Trypoxylus) and territories (fang blennies, Meiacanthus) [1-3]. Chemicals, including sprays, ointments and injected venoms, are another defence system used by animals. As with morphological weapons, venom can serve multiple purposes, including to facilitate feeding, in predation, and in defence when attacked [4]. Although rare, several taxa use venom for agonistic intraspecific competition (e.g. ghost shrimp, Caprella spp.; sea anemones, Actinia equina; cone snails, Conidae; male platypus, Ornithorhynchus anatinus) [4-6]. Another group of venomous mammals are the nocturnal slow lorises (Nycticebus) [7]. Slow loris bites often result in dramatic diagnostic wounds characterised by necrotic gashes to the head and extremities. Although these bites are the major cause of death of lorises in captivity, the function of this aggressive behaviour has never been studied in the wild [7]. Here, through an 8-year study of wounding patterns, territorial behaviour, and agonistic encounters of a wild population of Javan slow lorises (Nycticebus javanicus), we provide strong evidence that venom is used differentially by both sexes to defend territories and mates. VIDEO ABSTRACT.