Conservation of the Eurasian beaver Castor fiber: an olfactory perspective

• 1 Chemical communication in mammals includes an array of specific behaviours that are often ignored in terms of their potential relevance to conservation. Often used during territorial or social interactions between animals, chemical communication can also be used as a tool in reintroduction programmes. Reintroductions still exhibit high failure rates and methods to improve success should be investigated. The Eurasian beaver Castor fiber has been widely reintroduced across Europe after its near extinction in the 19th century.
• 2 Using olfactory studies in the beaver, we aim to demonstrate how scent transfers a range of information about the sender which can be used to monitor social and territorial behaviour along with general well‐being. Scent manipulation can be used to reduce human–beaver conflicts, and aid reintroduction success through reducing stress and territorial conflicts, and by influencing dispersal and settlement.
• 3 Two species of beavers, the Eurasian beaver and the North American beaver Castor canadensis, occupy freshwater habitats throughout North America and in parts of South America, most of Europe and parts of Asia. Most of the reviewed literature concerns the wild Eurasian beaver, its chemical communication and conservation; however, captive studies and those addressing North American beavers are also included.
• 4 Chemical communication is advanced and has been well documented in this highly territorial species. However, few studies directly link olfaction with conservation practices.
• 5 Olfactory studies in beavers can provide non‐invasive methods to monitor translocated animals and indicators of health. We conclude that chemical analysis, olfactory studies and behavioural manipulations involving semiochemicals have important impacts on conservation and can generate practical solutions to conservation problems including aiding animal capture, captive stress reduction, breeding pair formation and release site fidelity.

     

Chemosignalling of musth by individual wild African elephants (Loxodonta africana): implications for conservation and management

Elephants have extraordinary olfactory receptive equipment, yet this sensory system has been only minimally investigated in wild elephants. We present an in-depth study of urinary chemical signals emitted by individual, behaviourally characterized, wild male African elephants, investigating whether these compounds were the same, accentuated, or diminished in comparison with captive individuals. Remarkably, most emitted chemicals were similar in captive and wild elephants with an exception traced to drought-induced dietary cyanates among wild males. We observed developmental changes predominated by the transition from acids and esters emitted by young males to alcohols and ketones released by older males. We determined that the ketones (2-butanone, acetone and 2-pentanone, and 2-nonanone) were considerably elevated during early musth, musth and late musth, respectively, suggesting that males communicate their condition via these compounds. The similarity to compounds released during musth by Asian male elephants that evoke conspecific bioresponses suggests the existence of species-free 'musth' signals. Our innovative techniques, which allow the recognition of precise sexual and musth states of individual elephants, can be helpful to managers of both wild and captive elephants. Such sampling may allow the more accurate categorization of the social and reproductive status of individual male elephants.     

Modeling population viability of captive elephants in Myanmar (Burma): implications for wild populations

Captive Asian elephants Elephas maximus , used as work animals, constitute up to 22–30% of remaining Asian elephants. Myanmar has the largest captive population worldwide (∼6000), maintained at this level for over a century. We used published demographic data to assess the viability of this captive population. We tested how this population can be self-sustained, how many elephants must be supplemented from the wild to maintain it, and what consequences live capture may have for Myanmar's wild population. Our results demonstrate that the current captive population is not self-sustaining because mortality is too high and birth rates are too low. Our models also suggest ∼100 elephants year⁻¹ have been captured in the wild to supplement the captive population. Such supplementation cannot be supported by a wild population of fewer than 4000 elephants. Given the most recent expert estimate of ∼2000 wild elephants remaining in Myanmar, a harvest of 100 elephants year⁻¹ could result in extinction of the wild population in 31 years. Continued live capture threatens the survival of wild and captive populations and must stop. In addition, captive breeding should be increased. These measures are essential to slow the decline and extinction of all of Myanmar's elephants.     

中国亚洲象取食植物种类统计与分析

2016年1月至2017年12月,分别采用样线法和跟踪调查法对中国境内分布的野生亚洲象和云南西双版纳亚洲象种源繁育与救助中心的10头圈养亚洲象放养时取食植物进行调查,经分类鉴定,共收集到亚洲象取食植物111种,分属29目、42科、77属,其中新增亚洲象取食植物57种,分属于21目、32科、44属。与文献中记载的中国亚洲象植物性食物进行了汇总,统计得到新的中国亚洲象植物性食物名录,共计32目、62科、162属、240种。多样性分析结果显示,G-F多样性指数为0.84,表明中国野生及圈养亚洲象取食的植物具有较高的多样性,这些植物涉及蕨类植物门、裸子植物门和被子植物门。其中,圈养亚洲象采食植物种类共84种,西双版纳境内圈养亚洲象以禾本科(Gramineae)的甘蔗(Saccharum spp.)、玉米(Zea mays)、马唐(Digitaria spp.)、象草(Pennisetum purpureum)、粽叶芦(Thysanolaena latifolia)等为主要食物,而国内其它地方(北京动物园、广州动物园和昆明动物园)圈养的亚洲象主要饲喂高粱属(Sorghum)的高粱(S.bicolor)或苏丹草(S.sudanense)等。因此,在将来开展野生亚洲象栖息地保护、恢复及食物源基地项目规划与建设时应规避农作物和经济作物,优选亚洲象喜食、速生、生物量大的土著物种,如野芭蕉、棕叶芦、董棕、构树、重阳木、中平树、马唐属及竹类等植物,按照不同季节进行套种,为亚洲象提供更多可口食物。对于圈养亚洲象,需要设计足够面积的食物源基地供放养,补充食物种类,增加运动量,增强体质,提高其生活质量。     

Estrous state influences on investigative,aggressive, and tail flicking behavior in captive female Asian elephants

Females of species that live in matrilineal hierarchies may compete for temporally limited resources, yet maintain social harmony to facilitate cohesion. The relative degree of aggressive and nonaggressive interactions may depend on the reproductive condition of sender and receiver. Individuals can benefit by clearly signaling and detecting reproductive condition. Asian elephants (Elephas maximus) live in social matrilineal herds. Females have long estrous cycles (14–16 weeks) composed of luteal (8–12 weeks) and follicular (4–8 weeks) phases. In this study, we observed the behavior of four captive Asian elephant females during multiple estrous cycles over 2 years. We evaluated whether investigative, aggressive, and tail flicking behaviors were related to reproductive condition. Investigative trunk tip contacts showed no distinct pattern by senders, but were more prevalent toward female elephants that were in their follicular compared with their luteal phase. The genital area was the most frequently contacted region and may release reproductively related chemosignals. Aggression did not differ significantly with estrus; however, rates of aggression were elevated when senders were approaching ovulation and receivers were in the luteal phase. Females in the follicular phase may honestly advertise their condition. Contacts by conspecifics may serve to assess condition and reduce aggression. A behavior termed “tail flicking” was performed mainly during the mid‐follicular phase when estrogen and luteinizing hormone levels are known to spike. Tail flicking may disperse chemical signals in urine or mucus as well as act as a tonic signal that could provide a means of anticipating forthcoming ovulation by elephants and also for human observers and caretakers. Zoo Biol 27:167–180, 2008. © 2008 Wiley‐Liss, Inc.     

Cheap talk when interests conflict

Most evolutionary analyses of animal communication suggest that low-cost signals can evolve only when both the signaller and the recipient rank outcomes in the same order. When there is a conflict of interest between sender and receiver, honest signals must be costly. However, recent work suggests that low-cost signals can be evolutionarily stable, even when the sender and the receiver rank outcomes in different orders, as long as the interest in achieving coordination is sufficiently great. In this paper, we extend this body of work by analysing a game theory model that shows that low-cost signals can evolve when there are conflicts of interest and no interest in coordination, as long as individuals interact repeatedly. We also present an empirical example indicating that female rhesus macaques, Macaca mulatta, use honest, low-cost, vocal signals to facilitate interactions when conflicts of interest exist. Copyright 2000 The Association for the Study of Animal Behaviour.     

Endocrine Correlates of Musth in Free-Ranging Asian Elephants (Elephas maximus) Determined by Non-Invasive Faecal Steroid Hormone Metabolite Measurements

The occurrence of musth, a period of elevated levels of androgens and heightened sexual activity, has been well documented for the male Asian elephant (Elephas maximus). However, the relationship between androgen-dependent musth and adrenocortical function in this species is unclear. The current study is the first assessment of testicular and adrenocortical function in free-ranging male Asian elephants by measuring levels of testosterone (androgen) and cortisol (glucocorticoid – a physiological indicator of stress) metabolites in faeces. During musth, males expectedly showed significant elevation in faecal testosterone metabolite levels. Interestingly, glucocorticoid metabolite concentrations remained unchanged between musth and non-musth periods. This observation is contrary to that observed with wild and captive African elephant bulls and captive Asian bull elephants. Our results show that musth may not necessarily represent a stressful condition in free-ranging male Asian elephants.     

Concentration‐dependent chemosensory responses towards pheromones are influenced by receiver attributes in Asian elephants

The physical structure of a signal is not sufficient to determine its meaning. For chemical signals between conspecifics, this concept is termed “pheromonal parsimony.” The function of a compound depends not only on its molecular structure but also on its context, which can include signal concentration and various receiver attributes. We sought to investigate the contextual flexibility of chemosensory responses through bioassays with Asian elephant (Elephas maximus) sex pheromones of various concentrations (frontalin, from males, and (Z)‐7‐dodecenyl acetate [Z7‐12:Ac], from females). We hypothesized that elephants would respond stronger to higher concentrations, especially towards the opposite‐sex pheromone, and that receiver age and sexual experience would modify responses. We examined responses of 73 captive elephants to both compounds. Pheromone concentration impacted the rate of chemosensory response, which was further modified by the sex, age and/or sexual experience of the receiver. Response rates increased with concentration for each compound across both sexes. Experience shaped male responses with older, physiologically primed males responding more often. The interaction between experience and age affected female response to frontalin, but not to Z7‐12:Ac. Furthermore, response thresholds were modified by sexual experience in most cases: experienced animals generally had lower thresholds than inexperienced animals. Elephants responded to each solution according to its perceived relevance, including concentration. These results also indicate that receiver attributes (e.g., sex, age and sexual experience) may modify seemingly fixed chemosensory responses and further emphasize the flexibility of vertebrate communication systems.     

THE EVOLUTION OF INDIVIDUAL VARIATION IN COMMUNICATION STRATEGIES

Communication is a process in which senders provide information via signals and receivers respond accordingly. This process relies on two coevolving conventions: a “sender code” that determines what kind of signal is to be sent given the sender's state; and a “receiver code” that determines the appropriate responses to different signal types. By means of a simple but generic model, we show that polymorphic sender and receiver strategies emerge naturally during the evolution of communication, and that the number of alternative strategies observed at equilibrium depends on the potential for error in signal production. Our model suggests that alternative communication strategies will evolve whenever senders possess imperfect information about their own quality or state, signals are costly, and genetic mechanisms allow for a correlation between sender and receiver behavior. These findings provide an explanation for recent reports of individual differences in communication strategies, and suggest that the amount of individual variation that can be expected in communication systems depends on the type of information being conveyed. Our model also suggests a link between communication and the evolution of animal personalities, which is that individual differences in the production and interpretation of signals can result in consistent differences in behavior.     

Elephant Genomes Reveal Accelerated Evolution in Mechanisms Underlying Disease Defenses

Disease susceptibility and resistance are important factors for the conservation of endangered species, including elephants. We analyzed pathology data from 26 zoos and report that Asian elephants have increased neoplasia and malignancy prevalence compared with African bush elephants. This is consistent with observed higher susceptibility to tuberculosis and elephant endotheliotropic herpesvirus (EEHV) in Asian elephants. To investigate genetic mechanisms underlying disease resistance, including differential responses between species, among other elephant traits, we sequenced multiple elephant genomes. We report a draft assembly for an Asian elephant, and defined 862 and 1,017 conserved potential regulatory elements in Asian and African bush elephants, respectively. In the genomes of both elephant species, conserved elements were significantly enriched with genes differentially expressed between the species. In Asian elephants, these putative regulatory regions were involved in immunity pathways including tumor-necrosis factor, which plays an important role in EEHV response. Genomic sequences of African bush, forest, and Asian elephant genomes revealed extensive sequence conservation at TP53 retrogene loci across three species, which may be related to TP53 functionality in elephant cancer resistance. Positive selection scans revealed outlier genes related to additional elephant traits. Our study suggests that gene regulation plays an important role in the differential inflammatory response of Asian and African elephants, leading to increased infectious disease and cancer susceptibility in Asian elephants. These genomic discoveries can inform future functional and translational studies aimed at identifying effective treatment approaches for ill elephants, which may improve conservation.     

元江-李仙江流域亚洲象生境适宜性评价——基于荟萃分析和遥感大数据分析

生境适宜性评价是物种保护和生境管理与规划的基础。近几十年来,云南境内野生亚洲象数量剧增,外扩迁移事件频发,而新迁入区域生境适宜状况因物种出现点数据缺乏而难以评价,掣肘迁移亚洲象保护与风险防范应急。以亚洲象新近迁入的元江-李仙江流域为案例区,采用荟萃分析统计亚洲象生境评价因子,结合相关性分析和方差膨胀因子独立性检验,筛选出生境评价因子;基于开源遥感数据产品量化生境因子,综合主成分分析和层次分析计算生境评价因子权重,采用生境适宜性指数(Habitat Suitability Index, HSI)模型评价元江-李仙江流域亚洲象生境适宜性,并分析其景观格局。结果表明：(1)元江-李仙江流域亚洲象生境适宜性空间格局主要表现为由下游至上游呈递减趋势,最适生境主要分布于流域下游段,而流域上游段适宜生境少;(2)元江流域生境适宜性低于李仙江流域,且其生境斑块连接度更低、破碎化更严重;(3)2021年“北移象群”北迁沿程生境适宜性由西南向东北呈下降趋势。基于亚洲象生境适宜性评价结果,科学引导野生亚洲象迁入适宜生境区,以规避人象冲突,保障外迁亚洲象群及其活动区居民生命财产安全,服务于区域生物多样性保护与...     

Visual Cues Given by Humans Are Not Sufficient for Asian Elephants (Elephas maximus) to Find Hidden Food

Recent research suggests that domesticated species – due to artificial selection by humans for specific, preferred behavioral traits – are better than wild animals at responding to visual cues given by humans about the location of hidden food. \Although this seems to be supported by studies on a range of domesticated (including dogs, goats and horses) and wild (including wolves and chimpanzees) animals, there is also evidence that exposure to humans positively influences the ability of both wild and domesticated animals to follow these same cues. Here, we test the performance of Asian elephants (Elephas maximus) on an object choice task that provides them with visual-only cues given by humans about the location of hidden food. Captive elephants are interesting candidates for investigating how both domestication and human exposure may impact cue-following as they represent a non-domesticated species with almost constant human interaction. As a group, the elephants (n = 7) in our study were unable to follow pointing, body orientation or a combination of both as honest signals of food location. They were, however, able to follow vocal commands with which they were already familiar in a novel context, suggesting the elephants are able to follow cues if they are sufficiently salient. Although the elephants’ inability to follow the visual cues provides partial support for the domestication hypothesis, an alternative explanation is that elephants may rely more heavily on other sensory modalities, specifically olfaction and audition. Further research will be needed to rule out this alternative explanation.     

Pheromonal control: reconciling physiological mechanism with signalling theory

Pheromones are intraspecific chemical signals. They can have profound effects on the behaviour and/or physiology of the receiver, and it is still common to hear pheromones described as controlling of the behaviour of the receiver. The discussion of pheromonal control arose initially from a close association between hormones and pheromones in the comparative physiological literature, but the concept of a controlling pheromone is at odds with contemporary signal evolution theory, which predicts that a manipulative pheromonal signal negatively affecting the receiver's fitness should not be stable over evolutionary time. Here we discuss the meaning of pheromonal control, and the ecological circumstances by which it might be supported. We argue that in discussing pheromonal control it is important to differentiate between control applied to the effects of a pheromone on a receiver's physiology (proximate control), and control applied to the effects of a pheromone on a receiver's fitness (ultimate control). Critically, a pheromone signal affecting change in the receiver's behaviour or physiology need not necessarily manipulate the fitness of a receiver. In cases where pheromonal signalling does lead to a reduction in the fitness of the receiver, the signalling system would be stable if the pheromone were an honest signal of a social environment that disadvantages the receiver, and the physiological and behavioural changes observed in the receiver were an adaptive response to the new social circumstances communicated by the pheromone.     

亚洲象与竹/蕉分布隔离的生态效果及其保护对策探讨

把分布在滇南的西双版纳自然保护区内的亚洲象频繁出走 ,毁坏农作物、伤害人畜的原因与自然保护区片断化以及它们与其重要采食植物黄竹、野芭蕉等分布的隔离联系起来探讨 ,由此提出了对亚洲象保护的一些措施。由于亚洲象在西双版纳几乎没有天敌 ,而且也得到较好的保护 ,其群居自 1 976年以来 ,由 1 0 1头发展至约 2 5 0头 ,自然保护区片断化和食物匮缺成为亚洲象走出自然保护区的重要原因。相反 ,在没有亚洲象控制的地区 ,黄竹和野芭蕉等以其高效的繁殖能力以及耐火烧的特性而在一些刀耕火种的弃荒地和退化生态系统迅速发展 ,成为以它们各自为优势的群落 (丛 ) ,大幅度地降低了群落的物种多样性     

Birth seasonality and calf mortality in a large population of Asian elephants

In seasonal environments, many species concentrate their reproduction in the time of year most likely to maximize offspring survival. Asian elephants (Elephas maximus) inhabit regions with seasonal climate, but females can still experience 16‐week reproductive cycles throughout the year. Whether female elephants nevertheless concentrate births on periods with maximum offspring survival prospects remains unknown. We investigated the seasonal timing of births, and effects of birth month on short‐ and long‐term mortality of Asian elephants, using a unique demographic data set of 2350 semicaptive, longitudinally monitored logging elephants from Myanmar experiencing seasonal variation in both workload and environmental conditions. Our results show variation in birth rate across the year, with 41% of births occurring between December and March. This corresponds to the cool, dry period and the beginning of the hot season, and to conceptions occurring during the resting, nonlogging period between February and June. Giving birth during the peak December to March period improves offspring survival, as the odds for survival between age 1 and 5 years are 44% higher for individuals born during the high birth rate period than those conceived during working months. Our results suggest that seasonal conditions, most likely maternal workload and/or climate, limit conception rate and calf survival in this population through effects on maternal stress, estrus cycles, or access to mates. This has implications for improving the birth rate and infant survival in captive populations by limiting workload of females of reproductive age. As working populations are currently unsustainable and supplemented through the capture of wild elephants, it is imperative to the conservation of Asian elephants to understand and alleviate the effects of seasonal conditions on vital rates in the working population in order to reduce the pressure for further capture from the wild.     

Causes and correlates of calf mortality in captive Asian elephants (Elephas maximus)

Juvenile mortality is a key factor influencing population growth rate in density-independent, predation-free, well-managed captive populations. Currently at least a quarter of all Asian elephants live in captivity, but both the wild and captive populations are unsustainable with the present fertility and calf mortality rates. Despite the need for detailed data on calf mortality to manage effectively populations and to minimize the need for capture from the wild, very little is known of the causes and correlates of calf mortality in Asian elephants. Here we use the world''s largest multigenerational demographic dataset on a semi-captive population of Asian elephants compiled from timber camps in Myanmar to investigate the survival of calves (n = 1020) to age five born to captive-born mothers (n = 391) between 1960 and 1999. Mortality risk varied significantly across different ages and was higher for males at any age. Maternal reproductive history was associated with large differences in both stillbirth and liveborn mortality risk: first-time mothers had a higher risk of calf loss as did mothers producing another calf soon (<3.7 years) after a previous birth, and when giving birth at older age. Stillbirth (4%) and pre-weaning mortality (25.6%) were considerably lower than those reported for zoo elephants and used in published population viability analyses. A large proportion of deaths were caused by accidents and lack of maternal milk/calf weakness which both might be partly preventable by supplementary feeding of mothers and calves and work reduction of high-risk mothers. Our results on Myanmar timber elephants with an extensive keeping system provide an important comparison to compromised survivorship reported in zoo elephants. They have implications for improving captive working elephant management systems in range countries and for refining population viability analyses with realistic parameter values in order to predict future population size of the Asian elephant.     

社群压力对圈养雌性亚洲象动情的影响

亚洲象是中国一级濒危保护动物,改善圈养亚洲象的繁殖状况是亚洲象保护工作的重要内容之一。本研究旨在通过分析社群压力对圈养雌性亚洲象应激和动情状况的影响,从而探究圈养雌性亚洲象长期不动情的原因。通过放射性免疫法检测雌性亚洲象尿液中皮质醇、孕酮和雌二醇水平以及粪便中皮质醇和雌二醇水平,以雌激素水平反映亚洲象的动情状况,以皮质醇水平反映亚洲象的应激状态,以争斗行为和友好行为的发生情况反映圈养群体的社群压力。研究发现：(1)处于优势地位的个体表现出更多的斗争行为,处于从属地位的个体受到更多的攻击行为;(2)优势个体的皮质醇水平显著高于处于从属地位个体的皮质醇水平;(3)优势个体的雌二醇和孕酮具有小幅周期性波动,而从属个体的雌二醇和孕酮水平没有表现出明显的波动。从本研究结果可以看出圈养亚洲象群体中优势个体对从属个体的行为抑制可能是导致从属个体雌激素水平低,长期不能参与繁殖的原因。     

Oestrous state dynamics in chemical communication by captive female Asian elephants

In many mammals, reproductive status is revealed through chemical cues in urine. The reproductive status of receivers may influence their interest in such signals. For social mammals that live in matrilineal groups, females may benefit by detecting the reproductive condition of herdmates. Responses to urine during oestrous cycles of senders and receivers are potential indicators of signal functions. We examined the chemosensory responses, first by four captive female Asian elephants,Elephas maximus , over their oestrous cycles to familiar follicular and luteal phase urine and second by 14 different female Asian elephants to unfamiliar conspecific follicular and luteal phase urine. We asked whether females could distinguish the reproductive state of another female as measured by their differential response to luteal- and follicular-phase urine. We further examined the influence of the receiver's reproductive status on response levels. Females responded more with specific tactolfactory trunk behaviours to follicular- than to luteal-phase urine, but only when the receiving female was in her follicular phase. Like their male conspecifics, Asian elephant females can detect changes in the reproductive state of conspecifics. The functional significance of this ability has yet to be determined but may be related more to the resource holding power of females in follicular phase than to a means for females to synchronize oestrous cycles. Such female-female communication may have important effects on social group dynamics. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Prediction of estrus cyclicity in Asian elephants (Elephas maximus) through estimation of fecal progesterone metabolite: development of an enzyme-linked immuno-sorbent assay

Asian elephants (Elephas maximus), prominent “flagship species”, are listed under the category of endangered species (EN - A2c, ver. 3.1; IUCN Red List 2009) and there is a need for their conservation. This requires understanding demographic and reproductive dynamics of the species. Monitoring reproductive status of any species is traditionally being carried out through invasive blood sampling and this is restrictive for large animals such as wild or semi-captive elephants due to legal, ethical, and practical reasons. Hence, there is a need for a non-invasive technique to assess reproductive cyclicity profiles of elephants, which will help in the species’ conservation strategies. In this study, we developed an indirect competitive enzyme linked immuno-sorbent assay (ELISA) to estimate the concentration of one of the progesterone-metabolites i.e., allopregnanolone (5α-P-3OH) in fecal samples of Asian elephants. We validated the assay which had a sensitivity of 0.25 μM at 90% binding with an EC₅₀ value of 1.37 μM. Using female elephants, kept under semi-captive conditions in the forest camps of Mudumalai Wildlife Sanctuary, Tamil Nadu and Bandipur National Park, Karnataka, India, we measured fecal progesterone-metabolite (5α-P-3OH) concentrations in six animals and showed their clear correlation with those of serum progesterone, measured by a standard radio-immuno assay. Statistical analyses using a Linear Mixed Effect model showed a positive correlation (P < 0.1) between the profiles of fecal 5α-P-3OH (range: 0.5-10 μg/g) and serum progesterone (range: 0.1-1.8 ng/mL). Therefore, our studies show, for the first time, that the fecal progesterone-metabolite assay could be exploited to predict estrus cyclicity and to potentially assess the reproductive status of captive and free-ranging female Asian elephants, thereby helping to plan their breeding strategy.     

Primate Communication: By Nature Honest, or by Experience Wise?

We review evolutionary views on honesty and deception and their application to studies of nonhuman primate communication. There is evidence that some primate signals are likely to be accurate on the basis of costliness. They appear most often in contexts that include overtly competitive interactions in which unrelated individuals have limited access to information about one another. However, both game theoretic models and most empirical work suggest that costly signals are not often likely to be the basis for honest communication in nonhuman primates. Inexpensive signaling can exist in contexts wherein communication occurs among related animals, something common among many nonhuman primate societies. Another condition in which inexpensive signaling is possible and that is also typical of nonhuman primates, is when sender and receiver both benefit from coordinated interactions. Additionally, when individuals interact repeatedly and can use past interactions to assess the honesty of signals and to modify future response to signals, low-cost signals can evolve. Nonhuman primates appear to deal with the problem of deception via skeptical responding, which can be largely accounted for by learning rules and the fact that they live in stable social groups and can recognize one another and recall past interactions.