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The cestode Hymenolepis nana is a common parasite of humans and mice. Fecal shedding in the absence of clinical disease has previously been reported in ring-tailed lemurs ( Lemur catta). This report describes fatal, disseminated H. nana cestodiasis infection in an aged ring-tailed lemur in a zoological collection. The parasites were associated with severe multifocal to coalescing and regionally extensive pyogranulomatous hepatitis and moderate multifocal pneumonia. The morphology of the parasites was highly unusual. Profiles were variably sized, ellipsoid to irregularly serpiginous, lined by a thin tegument, and filled with lightly eosinophilic fibrillar stroma and numerous, round basophilic cells. Polymerase chain reaction targeting a portion of the 18S rRNA gene and DNA sequencing of the amplicon showed 100% homology with H. nana. Esther E. V. Crouch et al, Fatal Hymenolepis nana cestodiasis in a ring-tailed lemur (Lemur catta), Veterinary Pathology (, ) pp. . Copyright © 2021. DOI: 10.1177/03009858211042580. Users who receive access to an article through a repository are reminded that the article is protected by copyright and reuse is restricted to non-commercial and no derivative uses. Users may also download and save a local copy of an article accessed in an institutional repository for the user's personal reference. For permission to reuse an article, please follow our Process for Requesting Permission. Deposited by shareyourpaper.org and openaccessbutton.org. We've taken reasonable steps to ensure this content doesn't violate copyright. However, if you think it does you can request a takedown by emailing help@openaccessbutton.org.

Environmental enrichment is widely used to improve the quality of life of animals under human care. To successfully implement enrichment programs, it is important to evaluate their effect in different enclosure types since housing conditions may change depending on external factors, such as husbandry, management, or seasonal variation. This study investigates how ring-tailed lemur (Lemur catta) behaviour changes with the availability of enrichment items and the type of enclosure the animals are housed in. Through observations, we compared the behaviour of the lemurs in an indoor and outdoor enclosure, both without and with enrichment items. Although we observed enrichment effects, we found that enclosure type had a bigger effect on the lemurs’ behaviour. Additionally, behavioural changes induced by enrichment items differed between indoor and outdoor enclosures. These results indicate that the effectiveness of enrichment items may depend on the enclosure in which they are provided and consequently suggest that the impact of these programs should not be generalised over enclosure types. This highlights that the evaluation of environmental enrichment programs remains important when optimising zoo animal welfare.

During routine surveys of owl distributions on St. Catherines Island, we observed naturalized ring-tailed lemurs displaying antipredator responses to owl auditory cues. In the 32 y since the introduction of ring-tailed lemurs to the island, two successful depredation events by two different owl species have been documented. We investigated the behavioral response of ring-tailed lemurs to determine if they responded consistently to social calls from all three owl species present on St. Catherines Island despite size differences among the owl species that could affect the likelihood of them serving as predators on ring-tailed lemurs. We observed while ring-tailed lemurs responded to all the auditory owl calls, they exhibited more intense, longer and more consistent responses to the two larger owl species – the barred owl and great horned owl – relative to the small Eastern screech owl. These data suggest naturalized species are capable of learning threat-sensitive antipredator behaviors to novel predator communities.

High throughput sequencing of gut microbiota helps to understand the nutrition and health of the host. In this sense, it is an important tool for improving wildlife conservation and management. Little is known about the microbial communities occupying the gastrointestinal tract and the differences of microbiota among primate species. We conducted a microbiome study of the gastrointestinal tract in golden snub-nosed monkey (Rhinopithecus roxellanae), green monkey (Chlorocebus aethiops sabaeus), and ring-tailed lemur (Lemur catta), using the 16S rRNA gene amplicon sequencing and shotgun metagenomic sequencing. The gastrointestinal tract of all three primates were characterized by Firmicutes, Bacteroidetes, and Proteobacteria, although the relative abundance of these taxa differed across non-human primate species and gastrointestinal segments. In addition, the stomach of the golden monkey demonstrated a strong signal of microbial metabolic activity, especially in metabolizing carbohydrate. Prevotella multisaccharivorax and Selenomonas bovis were the main contributors to the function, and the enzymes involved in these metabolic activities were mainly derived from the glycoside hydrolase (GH) family. Microbial composition and function vary widely across non-human primate species and gastrointestinal segments. Therefore, while faecal samples are necessary for studying many mammals, particularly in the wild, data from the gastrointestinal tract have the potential to greatly improve our knowledge of host-gut microbe interactions. This information can contribute to the development of conservation interventions and animal care protocols in the wild and captivity to more fully describe a species’ nutritional needs and advance health and survival.

Several hypotheses have been offered to explain handedness in primates. In prosimian primates, including lemurs, left-hand orientation has been demonstrated; however, few studies have examined hand vs. mouth orientation. We compared preference for use of hand vs. mouth, single hand over both hands, and right hand vs. left hand in captive ring-tailed lemurs (Lemur catta), black-and-white ruffed lemurs (Varecia variegata), and collared brown lemurs (Eulemur collaris). For a male and female of each species, plus twin female offspring of ring-tailed lemurs, we made 100 observations per individual, recording hand vs. mouth use every 30 seconds as individuals performed enrichment and routine daily activities. Overall, these lemurs tended to be more handed than mouth-oriented, preferred to use a single hand more than both hands, and used their left hand more than their right. Males were more left-handed than females. Analysis of specific activities showed that eating was hand-oriented, while grooming was mouth-oriented. Results are also reported by species. Research on lemur use of hands vs. mouth could provide insight into the evolution of handedness in primates.

Ring-tailed lemurs reside in many animal collections worldwide. Lemur welfare may be a cause of concern due to some captive individuals exhibiting stereotypic behavior. Despite these concerns, there has been little exploration of methods of environmental enrichment for ring-tailed lemurs. Olfactory stimulation can enhance captive animal welfare by encouraging species-typical behaviors, enhancing behavioral diversity, and decreasing stereotypic behaviors. We aimed to investigate the effects of olfactory stimulation via lavender, peppermint, coconut, and prey odor upon the behavior of eight captive ring-tailed lemurs. We exposed the lemurs to six individual odor conditions (odor control, novel object control, lavender, peppermint, coconut, and Morio worms) and observed them for 4 hr a day for 3 days with an intervening period of 4 days between conditions. We recorded the lemurs' behavior under each condition using instantaneous scan sampling. We found significant effects of olfactory stimulation on the ring-tailed lemurs' behavior in the initial analysis but these did not survive correction for multiple testing. Overall, while our findings are suggestive of a general effect of olfactory stimulation on the captive ring-tailed lemurs they did not indicate a marked influence of olfactory condition. However, further investigation with a larger sample size and more biologically relevant odors may be beneficial to fully examine potential effects of olfactory stimulation in captive lemurs.

Responses to stress are unavoidable, adaptive mechanisms in humans and non-human animals. However, in humans, chronic stress has been linked to poor health outcomes and early mortality. Allostatic load, the physiologic dysregulation that occurs when an organism is exposed to chronic stressors, has been used to assess stress in humans, less work has been done using non-human primates. Our aim was to determine the relationship between allostatic load in ring-tailed lemurs (Lemur catta) under human care and potentially stressful individual, social, medical and husbandry factors, as well a sex and age. An allostatic load index (ALI) was calculated for 38 lemurs using six biomarkers measured in serum (albumin, cortisol, dehydroepiandrosterone-sulfate, DNA damage, glucose and prostaglandin E2). Potentially stressful factors were recorded over the lifetime of each lemur using medical and husbandry records. Animals with a higher percentage of time spent indoors, those kept in smaller average group sizes, and those with fewer minor group composition changes had, or tended to have, higher ALI. There was no relationship between ALI and sex or age. Some social and husbandry factors were associated with allostatic load in lemurs, indicating that this index may be a useful tool in assessing and determining factors contributing to stress of lemurs and other animals under human care.

We report on extremely rare events of lethal aggression in a semi-captive group of ring-tailed lemurs (Lemur catta) during the birth season 2014. This aggression was targeted against the two highest-ranking females. It led to their eviction from the group and following a final aggressive interaction four days later to their deaths caused by almost all the females and probably even two males of their former group. These events of targeted aggression erupted initially following an infanticide by the highest-ranking female directed at the offspring of a subordinate. Even for ring-tailed lemurs this is a very special case of changeover of power between two different matrilines in a group. In accordance with other studies we suggest that these events of targeted aggression were based on the growing group size and were acts of female reproductive competition during birth season. With the intensity of the events we add new aspects to the existing knowledge of aggressive interactions in ring-tailed lemurs.

1 In group‐living species with male dominance hierarchies where receptive periods of females do not overlap, high male reproductive skew would be predicted. However, the existence of female multiple mating and alternative male mating strategies can call into question single‐male monopolization of paternity in groups. Ring‐tailed lemurs (Lemur catta) are seasonally breeding primates that live in multi‐male, multi‐female groups. Although established groups show male dominance hierarchies, male dominance relationships can break down during mating periods. In addition, females are the dominant sex and mate with multiple males during estrus, including group residents, and extra‐group males—posing the question of whether there is high or low male paternity skew in groups. In this study, we analyzed paternity in a population of wild L. catta from the Bezà Mahafaly Special Reserve in southwestern Madagascar. Paternity was determined with 80–95% confidence for 39 offspring born to nine different groups. We calculated male reproductive skew indices for six groups, and our results showed a range of values corresponding to both high and low reproductive skew. Between 21% and 33% of offspring (3 of 14 or three of nine, counting paternity assignments at the 80% or 95% confidence levels, respectively) were sired by extra‐troop males. Males siring offspring within the same group during the same year appear to be unrelated. Our study provides evidence of varying male reproductive skew in different L. catta groups. A single male may monopolize paternity across one or more years, while in other groups, >1 male can sire offspring within the same group, even within a single year. Extra‐group mating is a viable strategy that can result in extra‐group paternity for L. catta males.

Assessing manual lateralization in non-human primates could be an optimal way to understand the adaptive value of this asymmetry in humans. Though many studies have investigated hand preferences in Old and New World monkeys and apes, fewer studies have considered manual lateralization in strepsirrhines, especially in experimental tasks. This study investigated hand preferences for a unimanual and a bimanual task of 17 captive ring-tailed lemurs (Lemur catta), housed at Parco Natura Viva (VR), Italy. The effect of age on handedness has been also investigated. The lemurs were tested on a unimanual task, in which subjects were restricted to using one hand to retrieve the food inside an apparatus, and on a bimanual task, in which lemurs had to use one hand to keep the apparatus door open while reaching with the other hand to retrieve the food inside it. At the population-level, our results revealed an asymmetrical hand use distribution, in particular a bias toward a right hand preference for food reaching in both the unimanual and the bimanual tasks. Furthermore, at the individual-level, the bimanual task seems to elicit a greater hand preference than the unimanual task. Results of this study underline the importance of experimental tasks in determining hand preference in strepsirrhines. Furthermore, as bimanual tasks elicited a stronger degree of lateralization, they appear to be more suited to investigate manual laterality. Finally, findings from this study highlight the presence of a right hand preference in ring-tailed lemurs, shedding new light on the evolution of human right handedness. Am. J. Primatol. 78:851-860, 2016. © 2016 Wiley Periodicals, Inc.

Objective To determine the minimum alveolar concentration (MAC) of sevoflurane for ring-tailed lemurs ( Lemur catta ) and aye-ayes ( Daubentonia madagascariensis ). Study design Prospective experimental trial. Animals Six adult ring-tailed lemurs, aged 1.3-11.2 years (median age: 8.26) and weighing a mean ± standard deviation (SD) of 2283 ± 254 g. Five adult aye-ayes, aged 4.4-19.3 years (median age: 8.0) and weighing 2712 ± 191 g. Methods Minimum alveolar concentration of sevoflurane was determined using a tail-clamp stimulus. The end-tidal sevoflurane ( Fe ’Sevo) concentration was increased or decreased by approximately 10% after a positive or negative response to tail clamping, respectively. This procedure was repeated until a positive and negative result were seen on two consecutive trials (i.e. a negative result was achieved and a single 10% decrease in Fe ’Sevo concentration resulted in a positive test). The MAC for that animal was determined to be the mean of the concentrations at the two consecutive trials. Results The mean ± SD MAC of sevoflurane for ring-tailed lemurs was 3.48 ± 0.55% and 1.84 ± 0.17 for aye-ayes. This represents a 47.1% higher MAC in ring-tailed lemurs compared to aye-ayes. Conclusions and clinical relevance The sevoflurane MAC was significantly higher in ring-tailed lemurs, compared to aye-ayes. The MAC of sevoflurane in aye-ayes is consistent with reported MAC values in other species. Extrapolation of sevoflurane anesthetic dose between different species of lemurs could lead to significant errors in anesthetic dosing.

Part of the conservation strategy of zoos is participation in ex situ conservation efforts in the form of captive breeding programs. Standardizing methods to describe and quantify behavior of animals housed at different institutions is an essential tool for understanding intra-species behaviors [Carlstead 2000; Carlstead 2002]. The primary objective of this study was to determine if a significant difference exists between the behavior of captive and wild populations of Lemur catta (ring-tailed lemurs) and to explore the implications of the results for captive management and reintroduction programs. Captive lemurs were found to be more inactive than wild lemurs and the type of enclosure (indoor or outdoor) had an impact on their species-typical sunning behavior. Zoos have several options to promote and maintain species-specific behaviors in captive populations.

The lifespan and age-specific fecundity of female ring-tailed lemurs (Lemur catta) were estimated from a 24-year longitudinal dataset based on individual identification at Berenty Reserve, Madagascar. The mean lifespan of females in 10-year (1989-1998) birth cohorts was 4.9 ± 4.9 years (n = 77), and the longest recorded lifespan in the population was 20 years. The mortality rate of adult females increased to ≥20% at 10-11 years old and reached 33-50% at 12-15 years old. Although the birth rate of old females (12-17 years old) was 72.0%, slightly lower than that of prime adult females (4-11 years old), i.e. 80.2%, no significant difference was found between them. Half of the females who reached the age of 12 years gave birth in the last year of their life. The oldest mother to give birth was 17 years old. These results suggest that most females can maintain reproductive performance in their later life and that there is no evidence for a postreproductive lifespan in this species.

In order to better understand the evolution of cognitive abilities in primates, information on cognitive traits of the most basal living primates can provide important comparative baseline data. Compared to haplorhine primates, lemurs have relatively smaller brains and reduced abilities to solve problems in the technical and social domain. However, recent studies have suggested that some cognitive abilities of lemurs are qualitatively equal to those of haplorhines. Here, we review studies investigating cognitive abilities in the technical and social domain of ring-tailed lemur cognition. In the physical domain, ring-tailed lemurs exhibit similar qualitative cognitive skills as other lemurs but also haplorhine primates. In the social domain, ring-tailed lemurs appear to be more skilled in visual perspective taking than other lemurs. Compared to other lemurs, they also have highly elaborated communicative skills. Moreover, within-group coalitions have been observed in female ring-tailed lemurs during rare events of female evictions but not in other lemur species. However, in several other aspects of social cognition, such as reconciliation and social learning, ring-tailed lemurs' cognitive abilities are equal to those of other lemurs. Thus, additional systematic comparative studies in physical and social cognition are required for a more comprehensive understanding of the processes of cognitive evolution among primates.

Polyspecific associations are well documented, but have rarely been observed in strepsirrhines. In this study we present a unique affiliative association between a female ring-tailed lemur (Lemur catta) and a group of southern bamboo lemurs (Hapalemur meridionalis) in south-east Madagascar. Our main research focused on H. meridionalis; however, due to the presence of the L. catta we treated her as a group member, including her in the focal sampling of Hapalemur social behaviour. We also recorded ad libitum data on all food species/items and any unique events or occurrences. Among observations, both species appeared to have a mutual understanding of vocalisations, behavioural synchronisation, dietary overlap, and possible service exchange, e.g., grooming. We also observed the L. catta occasionally attending to the bamboo lemur infant. This included grooming, baby-sitting, and even transporting the infant. The behavioural flexibility exhibited by both species has allowed the successful integration of the female ring-tailed lemur.

Bifidobacterium lemurum DSM 28807 T was isolated from the gastrointestinal tracts of ring-tailed lemurs ( Lemur catta ). Here, we report the first draft genome sequence of this organism.

Madagascar is known for its hypervariable climate with periodic droughts and cyclones, but little is known of the impact of such events on lemur physiology. We examined the effects of sequential weather periods, drought, normal, cyclone and post-cyclone, on hair cortisol concentrations (HCC) and body weight in wild ring-tailed lemurs, Lemur catta (n = 185), at the Beza Mahafaly Special Reserve in southwestern Madagascar. Data were modeled and analyzed by sex, age, and troop. Given the ecological consequences of extreme climatic perturbations, we hypothesized that drought and cyclone would significantly impact lemur HCC. Among adults, drought was associated with higher HCC than other periods and the lowest HCC was associated with the post-cyclone period. Adult females had greater variation in HCC during drought and males had greater variation during cyclone and Post-cyclone periods, suggesting sexes were differentially affected in terms of how individuals responded to extreme weather events. Low HCC in the post-cyclone period followed a 12-month period of reduced availability of primary and fallback food resources. Based on the known extreme and chronic nutritional stress during this time, our results indicate hypocortisolism in the animals included in our analysis. Higher HCC in sub-adults during the cyclone also suggests that immature lemurs may experience extreme weather events differently than adults. Body weight, used as a gauge for environmental stress, was lowest during the post-cyclone for sub-adults, young adults, and adults. Body weight did not differ by sex among adults across any of the weather events. Overall, ring-tailed lemur's HCC appear to be more immediately impacted by drought, or stressors associated with that specific weather event, and influenced by the long-term impact of cyclones on resource availability evidenced by data from the post-cyclone period.

Visual illusions are commonly used in animal cognition studies to compare visual perception among vertebrates. To date, researchers have focused their attention mainly on birds and mammals, especially apes and monkeys, but no study has investigated sensitivity to visual illusions in prosimians. Here we investigated whether lemurs (Lemur catta) perceive the Delboeuf illusion, a well-known illusion that occurs when subjects misperceive the relative size of an item because of its surrounding context. In particular, we adopted the spontaneous preference paradigm used in chimpanzees and observed lemurs’ ability to select the larger amount of food. In control trials, we presented two different amounts of food on two identical plates. In test trials, we presented equal food portion sizes on two plates differing in size: If lemurs were sensitive to the illusion, they were expected to select the food portion presented on the smaller plate. In control trials, they exhibited poor performance compared to other mammals previously observed, being able to discriminate between the two quantities only in the presence of a 0.47 ratio. This result prevented us from drawing any conclusion regarding the subjects’ susceptibility to the Delboeuf illusion. In test trials with the illusory pattern, however, the subjects’ choices did not differ from chance. Our data suggest that the present paradigm is not optimal for testing the perception of the Delboeuf illusion in lemurs and highlight the importance of using different methodological approaches to assess the perceptual mechanisms underlying size discrimination among vertebrates.

Ring-tailed lemurs (Lemur catta) at the Beza Mahafaly Special Reserve (BMSR), Madagascar display a high frequency of individuals with notable and sometimes extreme tooth wear. Adult lemurs display a range of tooth wear even among individuals of the same age, but we do not know at what age this variation first appears. This study's goal was to determine whether wear variation occurs in younger wild lemurs. Based on the decade-long study of ring-tailed lemur feeding and dental ecology at BMSR, we hypothesized that younger, natal lemurs (under 5 years of age), would display variation in their degree of tooth wear that would correspond to microhabitat differences, given differences in food availability in different troops' home ranges. We also hypothesized that wear would differ between sexes at this young age, given differences in feeding between males and females in this population. Hypotheses were tested using dental topographic analyses using dental impressions collected from known-aged lemurs across 10 years at BMSR. Results illustrate significant differences in wear-related tooth topography (i.e., relief and slope, presented here as "occlusal lift") for microhabitat, sex and troop affiliation among lemurs under 5 years of age in this population. Although, all lemurs in this population consume mechanically challenging tamarind fruit, those in more disturbed habitats eat additional introduced foods, some of which are also mechanically challenging. Thus, dietary variation is the likely cause of variation in tooth wear. The wear variation we show at a young age suggests caution when assigning age based on tooth wear in living and fossil primates. These wear-related tooth shape changes early in life, which reflects sex, habitat variation and levels of anthropogenic disturbance, may potentially impact reproductive fitness later in life.

Play signals are commonly used by animals to communicate their playful motivation and to limit the risk that rough acts are misunderstood by playmates. The relaxed open mouth is the most common facial expression performed during play in many mammals and represents the ritualized version of the movement anticipating a play bite. The signaling nature of this expression has been proven in many haplorrhine species but never demonstrated in strepsirrhines. Our purpose was assessing whether, also in strepsirrhines, the relaxed open mouth has an actual communicative function. We studied wild ring-tailed lemurs (Lemur catta), characterized by highly social habits including intense playful interactions. They largely use playful signals, mostly performed with the black and white tail. The signaling function of the tail (tail play) has been widely demonstrated. We analyzed both tail play and the relaxed open mouth to verify how their distribution is affected by different play variables (e.g., play session symmetry, number of play mates, previous use of the same pattern). Indeed, ring-tailed lemurs use the relaxed open mouth as a communicative signal during play. Relaxed open mouth was more frequent during unbalanced interactions showing the highest asymmetry in the patterns performed by the two players (offensive/neutral). Compared to tail play, relaxed open mouth was more frequent during dyadic than polyadic interactions and, as a highly directional signal, it was more frequently replicated by the play mate. Therefore, the relaxed open mouth needs to be performed face-to-face so that signal detection can be optimized. Similar to previous findings in monkeys and apes, the relaxed open mouth in lemurs seems to be a ritualized signal used to engage and, perhaps, sustain playful interaction.

The ring-tailed lemur (Lemur catta) is a quadruped arboreal primate primarily distributed in south and south-western Madagascar. This study was carried out to describe the normal radiographic thoracic anatomy of the ring-tailed lemur as a reference for clinical use.Radiography of the thorax was performed in 15 captive ring-tailed lemurs during their annual health examinations.Normal radiographic reference ranges for thoracic structures were established and ratios were calculated, such as the vertebral heart score (VHS). The mean VHS on the right lateral and dorsoventral views was 8.92 ± 0.47 and 9.42 ± 0.52, respectively.Differences exist in the normal radiographic thoracic anatomy of primates. Knowledge of the normal radiographic thoracic anatomy of individual species is important and fundamental to assist in clinical cases and for accurate diagnosis of diseases.

Monozygotic (MZ) twinning is generally considered to be rare in species other than human. We inspected sex ratios in European zoo-bred ring-tailed lemurs (Lemur catta), revealing a significant excess of same-sex twins. Of 94 pairs, 60 (64%) were either both males or both females (p= .004). Application of the Weinberg differential rule argues that 27% of all twins in this species are MZ pairs. In this protected species, where twinning is commonplace (~50% of newborns are twins), the probable existence of frequent MZ twinning has ramifications for breeding programs aimed to maximize genetic diversity, and suggests that twin studies in a species other than human could have potential as a medical research tool.

In mammals, purring has been described in mostly affiliative contexts. In the ring-tailed lemur (Lemur catta), both males and females purr, but only males were observed purring in agonistic contexts. In order to determine whether male ring-tailed lemurs purr as aggressive displays during intrasexual agonistic encounters, 480 h of focal data were collected on 25 adult males from Beza Mahafaly Special Reserve, Madagascar, from March to July 2010. The male purring rate increased during periods of male-male agonism when compared to times without intrasexual agonism, and the purring rate was positively correlated with male dominance rank. However, the purring rate was not significantly higher during winning agonistic interactions when compared with losing encounters. My results indicate that the male ring-tailed lemur purr is used most frequently as an agonistic vocalization in male-male encounters, in addition to being used less frequently in other social contexts, including during tail-waving at females, resting, scent-marking, feeding and copulation. Dominant males have higher purring rates across social situations, suggesting that the purring rate may be driven by intrinsic male qualities rather than functioning as a meaningful signal in each disparate social context. Male purring in intrasexual agonistic encounters can be added to previously described social contexts for ring-tailed lemur purring.

Long calls are sex-specific vocalizations used for mate attraction or mate defense in many animal species. Ring-tailed lemurs (Lemur catta), female-dominant strepsirrhines, have a male-specific long call termed a howl, with proposed functions that have never been empirically tested. I aimed to investigate why ring-tailed lemur males howl and to test whether the mate defense and mate attraction hypotheses for long-calling were applicable to this species. From March to July 2010, I collected 600 h of focal data on 25 males aged ≥3 year at Beza Mahafaly Special Reserve, Madagascar. I observed each male continuously for 30 min at a time and noted all agonism using one–zero sampling at 2.5-min intervals. I calculated male dominance rank from these data. I recorded days when female estrus occurred and noted howling and intergroup encounters using all-occurrences sampling. Howling rate was not significantly related to female estrus or male dominance rank, providing no support for the mate attraction hypothesis or the intragroup mate defense hypothesis. In contrast, the intergroup mate defense hypothesis was strongly supported. During intergroup encounters, male howling rate significantly increased compared to howling rate at times without other groups present, and a greater number of males participated in multimale howling choruses when compared to times without nongroup members present. My results suggest that male ring-tailed lemurs howl to advertise their presence and location to other groups, but not to male or female members of their own group. Howling could discourage male immigration by advertising the number of males already present in a group. Long calls are used for similar mate defense purposes during intergroup encounters by other primates, including Thomas langurs (Presbytis thomasi) and chacma baboons (Papio ursinus).

The effect of the zoo environment on captive animals is an increasingly studied area of zoo research, with visitor effects and exhibit design recognized as two of the factors that can contribute to animal welfare in captivity. It is known that in some situations, visitors may be stressful to zoo-housed primates, and this may be compounded by environmental factors such as the weather, the time of day, and zoo husbandry routines. Exhibit design and proximity of the public are also known to influence behavioral response of primates to visitors; however, there is minimal research on free-ranging zoo animals, even though they are potentially subjected to intense interactions with visitors. The current study explores the effect of the zoo environment, several visitor variables and specific animal-visitor interactions on the behavior of free-ranging ring-tailed lemurs (Lemur catta) at Fota Wildlife Park, Ireland. Data were obtained through scan samples collected over 18 months (n = 12,263) and analyzed using a range of statistical tests, including general estimating equations (GEE). Results demonstrate that the free-ranging lemurs' behavior at Fota Wildlife Park is affected by season, weather and time of day. Similarities in feeding behavior exist between the free-ranging group and lemurs in the wild when resources are plentiful. Visitor variables had a limited effect on lemur behavior and behavioral diversity level. Lemurs rarely reacted to visitors when specific interactions were considered. Generally, the results indicate that the ring-tailed lemurs in this study have adapted well to the zoo environment and habituated to visitors.

Lemurs are the most threatened group of mammals on earth. Lemur catta (ring-tailed lemur) represents one of the most iconic lemur species and faces numerous anthropogenic threats in the wild. In this study, we present population estimates from 32 sites across the range of L. catta, collected from primary and secondary data sources, to assess the number of ring-tailed lemurs left in the wild. We estimate that there are approximately 2,220 individual L. catta remaining in the 32 sites considered. We note local extinctions of populations of L. catta in at least 12 of the 32 sites examined, and that significantly more extinctions occurred in areas without some form of protection. This decrease in extant populations could represent a decrease of more than 95% of all ring-tailed lemurs in Madagascar since the year 2000. While these results should be considered preliminary, we stress the rapid decline of the species and note that habitat loss, bushmeat hunting and the illegal pet trade are driving populations to local extinction. Based on the data presented here, urgent and immediate funding and conservation action are crucial to ensure the viability of the remaining wild populations of ring-tailed lemurs.

Animals communicating via scent often deposit composite signals that incorporate odorants from multiple sources; however, the function of mixing chemical signals remains understudied. We tested both a ‘multiple-messages’ and a ‘fixative’ hypothesis of composite olfactory signalling, which, respectively, posit that mixing scents functions to increase information content or prolong signal longevity. Our subjects—adult, male ring-tailed lemurs (Lemur catta)—have a complex scent-marking repertoire, involving volatile antebrachial (A) secretions, deposited pure or after being mixed with a squalene-rich paste exuded from brachial (B) glands. Using behavioural bioassays, we examined recipient responses to odorants collected from conspecific strangers. We concurrently presented pure A, pure B and mixed A + B secretions, in fresh or decayed conditions. Lemurs preferentially responded to mixed over pure secretions, their interest increasing and shifting over time, from sniffing and countermarking fresh mixtures, to licking and countermarking decayed mixtures. Substituting synthetic squalene (S)—a well-known fixative—for B secretions did not replicate prior results: B secretions, which contain additional chemicals that probably encode salient information, were preferred over pure S. Whereas support for the ‘multiple-messages’ hypothesis underscores the unique contribution from each of an animal's various secretions, support for the ‘fixative’ hypothesis highlights the synergistic benefits of composite signals.

Metabolic costs of resting and locomotion have been used to gain novel insights into the behavioral ecology and evolution of a wide range of primates; however, most previous studies have not considered gait-specific effects. Here, metabolic costs of ring-tailed lemurs (Lemur catta) walking, cantering and galloping are used to test for gait-specific effects and a potential correspondence between costs and preferred speeds. Metabolic costs, including the net cost of locomotion (COL) and net cost of transport (COT), change as a curvilinear function of walking speed and (at least provisionally) as a linear function of cantering and galloping speeds. The baseline quantity used to calculate net costs had a significant effect on the magnitude of speed-specific estimates of COL and COT, especially for walking. This is because non-locomotor metabolism constitutes a substantial fraction (41-61%, on average) of gross metabolic rate at slow speeds. The slope-based estimate of the COT was 5.26 J kg(-1) m(-1) for all gaits and speeds, while the gait-specific estimates differed between walking (0.5 m s(-1) : 6.69 J kg(-1) m(-1) ) and cantering/galloping (2.0 m s(-1) : 5.61 J kg(-1) m(-1) ). During laboratory-based overground locomotion, ring-tailed lemurs preferred to walk at ~0.5 m s(-1) and canter/gallop at ~2.0 m s(-1) , with the preferred walking speed corresponding well to the COT minima. Compared with birds and other mammals, ring-tailed lemurs are relatively economical in walking, cantering, and galloping. These results support the view that energetic optima are an important movement criterion for locomotion in ring-tailed lemurs, and other terrestrial animals.

In September 2008, two ring-tailed lemurs (Lemur catta), comprising a mother–daughter pair, at the Greenville Zoo, Greenville, South Carolina, USA, were diagnosed with cuterebrid myiasis (Diptera: Oestridae) subsequent to sudden death of the adult lemur. On necropsy, a single bot warble was discovered in the subcutis of the axillary region. Histopathology revealed a severe eosinophilic bronchopneumonia. The juvenile lemur was inspected and found to have warbles on three separate dates in late September. One representative bot fly larva was identified as a Cuterebra sp. that normally infests lagomorphs in the southeastern United States. Cuterebrid myiasis is rarely reported in lemurs and has not been previously associated with pneumonia or death in these animals.

Telazol has been commonly used for field anesthesia of wild lemurs, including ring-tailed lemurs (Lemur catta). Telazol alone provides good induction, but doesn't cause adequate muscle relaxation and sedation for collecting consistent somatic measurements and high-quality dental impressions that are sometimes needed. Variability in induction response has been seen between individuals that have received similar dosages, with young lemurs seeming to need more anesthetic than mature lemurs. This investigation evaluated Telazol induction in young (2.0-4.9 yr) and mature (or = 5.0 yr) ring-tailed lemurs and compared postinduction supplementation with medetomidine or medetomidine-butorphanol. Forty-eight lemurs were anesthetized with Telazol administered via blow dart; then, 20 min after darting, they were supplemented via hand injection with either medetomidine (0.04 mg/ kg) or medetomidine-butorphanol (0.04 mg/kg and 0.2 mg/kg, respectively). The odds ratio for young lemurs to need more than one dart for induction, relative to mature lemurs, was 3.8, even though the initial dose of Telazol received by young lemurs (19 +/- 7 mg/kg) was significantly higher than the initial dose administered to mature lemurs (12 +/- 5 mg/kg). The total Telazol dosage was also significantly different between young lemurs (33 +/- 15 mg/kg) and mature lemurs (18 +/- 9 mg/kg). Both medetomidine and medetomidine-butorphanol provided good muscle relaxation and sedation for all procedures. Physiologic values were similar between the two protocols. Oxygen saturation by pulse oximetry was generally good, although there were a few SaO2 values90%. Recoveries were smooth, but long. Time to head up was correlated with total Telazol dosage in mature lemurs. In young lemurs, time to standing was correlated with Telazol induction dosage and time of last Telazol administration. Lemurs that received hand injections of Telazol took longer to recover than those that did not. Further refinements are needed to increase induction reliability and to decrease recovery time, particularly in young lemurs.

Lemur catta Linnaeus, 1758, is a lemurid commonly called the ring-tailed lemur. A strikingly colored, long-tailed lemur, it is the only species in the genus Lemur. It occurs in the southern half of Madagascar, where it inhabits a variety of forested habitats. It is listed as Vulnerable by the International Union for Conservation of Nature and Natural Resources.

Echinococcus multilocularis is a tapeworm responsible in its larval stage for alveolar echinococcosis, a disease which is lethal when left untreated. Multivesiculated parasitic lesions in the liver were diagnosed at necropsy in a captive-born nutria (Myocastor coypus) and in a ring-tailed lemur (Lemur catta) which had been in a French zoo for 16months. Molecular analyses confirmed the diagnosis of E. multilocularis obtained by histological analyses. These were the first cases of infection by E. multilocularis reported in lemurs in Europe, and the first case in nutria in European enclosures. Lemurs are confirmed to be particularly sensitive to E. multilocularis with a massive infection. In both cases, the infection appears to have been contracted in the zoo indirectly via environmental contamination by feces from roaming foxes. Due to the large endemic area for E. multilocularis, the increasing prevalence in foxes in France, and an increase in awareness of the disease, other cases of infection in captive animals will probably be recorded in France in the coming years.

Feeding enrichment is a commonly proposed way to promote natural behavior, greater activity levels and improved health in captive primates, but in many cases the methods have not been formally tested, especially for particular groups such as lemurs. We investigated whether simple changes in food presentation could increase activity levels in captive ring-tailed lemurs ( Lemur catta ) and reduce the removal by lemurs of food intended for hyrax ( Procavia capensis ) and porcupine ( Hystrix cristata ) (pirating) housed in the same enclosure. We varied two aspects of food presentation for a population of eight L. catta housed at the Santa Ana Zoo in a repeated measures, factorial design: the presence or absence of browse in food boxes, and the clumping or spatial separation of food boxes. The number of animals in the group engaged in different categories of behaviors was recorded with scans every 60 s for 2 h after feeding. Neither the addition of browse ( P = 0.58) nor spatial separation of food boxes ( P = 0.13) increased lemur activity levels in the first 70 min post-feeding, but addition of browse significantly raised activity in the last 50 min ( P = 0.038). Adding browse more than doubled (to 78%) the percentage of observation periods when at least one lemur was active after 70 min post-feeding and increased the percentage of active behaviors during that period from less than 4% to over 13%. Spatially separating food boxes reduced pirating ( P = 0.021), but addition of browse did not ( P = 0.89). Separating food boxes reduced pirating by nearly half. Given that these methods are inexpensive, easy to implement and also led to significant behavioral changes, simple feeding enrichment of this kind is a useful potential tool for captive ring-tailed lemur management.

In 1999, we measured the body mass of 101 wild ring-tailed lemurs (Lemur catta) inhabiting the Berenty Reserve, Madagascar. In addition, we counted the number of ticks [Haemaphysalis (Rhipistoma) lemuris Hoogstraal, 1953] infesting their facial skin and external auditory meatuses. For both males and females, the body mass appeared to increase until the age of 3 years. With the apparent exception of infants, there were no sexual differences in body mass. Within a group, higher-ranked adult males tended to be heavier than lower-ranked males. In contrast, there was no consistent correlation between the body mass of females and their ranks. Among the study groups, there was a small difference in body mass and significant difference in the number of ticks infesting the facial skin and external auditory meatuses. In particular, lemurs of a group who inhabited an area of gallery forest in the study area exhibited the smallest values of body mass and were severely infested with ticks. Such group variations were not consistently correlated with the reproductive parameters of the study groups. In three groups moderately infested with ticks, ticks infested adult males and subadults more heavily than adult females, juveniles, and infants.

Because ring-tailed lemurs (Lemur catta) show handedness during feeding, I investigated whether they also show lateral preference in the limbs used by males during scent marking. During a bout of marking, a male alternates a variable number of shoulder rubs and arm marks. It is a complex social signal that has both olfactory and visual components and is frequently part of intrasexual agonistic interactions. I used all-occurrences behavior sampling, manually recorded all totally visible bouts, and required a minimum of 50 bouts per animal. The subjects included six wild and four group-living captive male lemurs. I considered limb usage to be lateralized for each individual lemur if the frequency of use of one side was significantly above chance as determined by binomial z-scores at P

Adaptive explanations for social grouping assume that there are fitness consequences associated with group size, and individuals maintain membership in groups of favorable size to maximize fitness. Here I examine fecal cortisol concentrations as a hormonal measure of stress to assess the relative well-being of Lemur catta in groups of different size and in seasons of normal and low tamarind fruit availability. I test the hypotheses that there is an optimal group size at which cortisol is lowest and that optimal group size changes in food-scarce conditions. I collected 799 fecal samples from 87 individuals in seven free-ranging L. catta groups at Berenty Reserve, Madagascar, over a 1-year period (August 1999--July 2000) and determined fecal cortisol concentrations using a radioimmunoassay. Expressing these as residuals from monthly population means to control for temporal fluctuations in cortisol concentration, I calculated mean fecal cortisol levels for each animal in seasons of normal and low tamarind fruit abundance and over the entire year. Overall, females exhibited lowest mean cortisol levels in groups of intermediate size, suggesting that there are benefits to maintaining membership in these groups. Females in groups that were atypically large or small for their habitat type had higher mean cortisol levels than typical groups. Cortisol levels increased in food-scarce conditions for larger groups, suggesting that intergroup competitive advantages do not outweigh intragroup feeding competition at this time. Group size may be optimized for long-term average conditions, and short-term stresses may intermittently alter the costs associated with group size. Copyright 2005.

This research explores the effects of posture, sex, and living condition on hand and side preferences of semi-free-ranging, adult ring-tailed lemurs (Lemur catta) housed at the Duke University Primate Center in Durham, NC. Data were collected on 11 adult individuals (five females and six males) during normal daily activities over a ten-week period from May–July 2001. Variables analyzed in this study include unimanual behaviors (i.e., reach, hold, and limb used to start locomotion) and other potentially lateralized behaviors that do not involve handuse (i.e., whole-body turning and tail position). The data were analyzed to investigate potential individual and population level side biases for each behavior; potential sex biases in side preference for each behavior; and for ‘reach’, potential effects of posture (sitting, tripedal stance, or bipedal stance) on individual hand preferences. Additionally, to investigate potential effects of living condition on lateral biases, the data from this study were compared to data collected on the same individual Lemurs living under more restrictive living conditions during the previous year. Largely, as predicted based on available literature, we found that there was a significant sex difference across all hand-use categories and for whole-body turning, and that posture was a significant factor in the expression of hand preference for reaching. Contrary to previous research, the effect of living condition on lateral preferences was minimal, and no side preferences were found at the population level for any of the behaviors analyzed.

The subject of the present study was a tongue of the ring-tailed lemur (Lemur catta), which was provided by the Cooperation Research Program of the Primate Research Institute, Kyoto University. Using a scanning electron microscope, we observed the fine structures of the lingual papillae, which are distributed on or just under the surface of the body of the tongue of this animal. We compared our results with other primates and other mammalian species, and discussed the evolution of the papillae from the viewpoint of comparative anatomy.The filiform and fungiform papillae were observed on the dorsum of the tongue body in the ring-tailed lemur. The filiform papillae are arranged in a circular pattern chracteristic of the primates. Each circle consists of 13-18 cone-like papillae. There are some differences in size and shape of the papillae between the anterior and posterior parts of the tongue. The general morphology is similar both in the epithelial and connective tissue papillae. In contrast, the fungiform papillae show a hemispheric shape on the epithelium, while they are of a cactus-like shape in the connective tissue. This feature of the fungiform papillae is quite similar to those observed in several species of insectivores and lagomorphs.According to the present study, together with the results by previous authors, it appears that in lemurs, which are classified in prosimians, the filiform papillae show a morphology chracteristic of the order primates, but the fungiform papillae still retain a primitive feature relative to the Anthropoidea.

In ring-tailed lemurs, Lemur catta, the factors modulating hypothalamic-pituitary-adrenal (HPA) activity differ between wild and semi-free-ranging populations. Here we assess factors modulating HPA activity in ring-tailed lemurs housed in a third environment: the zoo. First we validate an enzyme immunoassay to quantify levels of glucocorticoid (GC) metabolites in the faeces of L. catta. We determine the nature of the female-female dominance hierarchies within each group by computing David's scores and examining these in relation to faecal GC (fGC). Relationships between female age and fGC are assessed to evaluate potential age-related confounds. The associations between fGC, numbers of males in a group and reproductive status are explored. Finally, we investigate the value of 7 behaviours in predicting levels of fGC. The study revealed stable linear dominance hierarchies in females within each group. The number of males in a social group together with reproductive status, but not age, influenced fGC. The 7 behavioural variables accounted for 68% of the variance in fGC. The amounts of time an animal spent locomoting and in the inside enclosure were both negative predictors of fGC. The study highlights the flexibility and adaptability of the HPA system in ring-tailed lemurs.

A case of fatal infection caused by larval forms of Taenia martis in a ring-tailed lemur (Lemur catta) living in the Rome zoological garden is described. The animal, living in a semi-natural pen with other 15 conspecific individuals and being fed with fresh fruit and vegetables, yoghurt and eggs, was transported to the Istituto Zooprofilattico of Rome for post-mortem examination. The anamnesis included, ten days before the death, apathy, lack of appetite, abdominal distension and diarrhoea. A severe exudative fibrinous-purulent peritonitis with numerous adhesions between the abdominal wall and the bowel loops was detected. After intestine removal, two free and viable, 4 cm long, whitish, leaf-like parasitic forms were pinpointed. Macroscopic examination of the two parasites allowed their identification as larval stages of cestodes, identified via molecular analysis as T. martis metacestodes. This report represents the first record of T. martis infection in the host species and in a zoological garden and for the pathological relevance of the infection.

In the Berenty Reserve, Southern Madagascar, of the 16 deliveries recorded for ring-tailed lemurs from 1995 and 1997, 10 cases occurred during daytime hours (07:00 – 16:59). Four mothers lost contact with their troop members during parturition, but were able to rejoin them after about one hour. No attack from the individuals of the neighboring groups or predators on the mothers or their newborn infants was observed, whereas, other females of the mother’s own troop attacked them after delivery in two cases. Such daytime delivery may have originated from the circadian rhythm of the species’ nocturnal ancestors, who might have given birth during the daytime. This pattern may now be a non-adaptive feature, in agreement with the “evolutionary disequilibrium hypothesis (EVDH).”

From 1989 to 1998, 204 live births were recorded for ring-tailed lemurs (Lemur catta) at Berenty, Madagascar. Excluding unknown birth dates, the peak month of birth was September, with 82.0% (146/178) occurring during this period. The offspring sex ratio (1∶1.19) was not significantly different from 1∶1, and there was no association with the mother's age. The first births occurred at the ages of 2 to 4 yr. The annual birth rate was very low at the age of 2 yr (11.1%), but increased thereafter: to 50.0% at the age of 3 yr, and to 75–85% at the age of 4 or more years. Multiple births were very rare, since only three sets of twins and one set of triplets were recorded. As for the interbirth interval, a one-year interval was the most common (92.2%). Infant mortality within the first year was 37.7% (77/204). Neonatal mortality within the first month accounted for 31.2% of all infant dealths.

A recent survey of the high-mountain zone of the Madagascar Parc National (PN) d'Andringitra revealed the presence of an apparently isolated troop of the ring-tailed lemur Lemur catta. These animals display phenotypic and ecological characteristics that are unusual for the monotypic genus Lemur, thus raising the possibility that they are members of a different undescribed species. We present analyses of two mitochondrial genes to test the hypothesis that L. catta from Andringitra should be considered a distinct species. The results indicate that taxonomic revision is not warranted under the expectations of the phylogenetic, coalescent, or biological species concepts. Rather, the genetic patterns observed among the Andringitra and lowland mitochondrial haplotypes are consistent with those expected for a single species.

To determine how the birth and development of infants influence their mothers' social relationships with other adult troop members, we observed two free-ranging troops of ring-tailed lemurs (Lemur catta) at the Berenty Reserve, Madagascar. The number of acts of affiliative contact that the mothers received from other adult troop members during the first and second months of infant life were significantly higher than those before they gave birth, and the values during the third month were as low as that before giving birth. Two mothers received acts of affiliative contact less frequently after their infants died, compared with the values while the infants were alive. On the other hand, more than 95% of all acts of licking of infants by adult troop members other than their mothers occurred when the infants were in contact with their mothers. These findings suggest that infants per se and mothers per se were not attractive, but rather the mother-infant pair was attractive to other troop members. Acts of infant-licking were observed in almost all mother-mother pairs and mother-non-mother adult female pairs, and in two thirds of mother-adult male pairs. Moreover, the frequency of infant-licking was not affected by female parity, female and male dominance rank, or infant sex. Therefore, acts of infant-licking, which are widespread among troop members, may function to make or maintain stable social relationships.

Feeding behavior in two groups of semi free-ranging primates, lion tailed macaques (LTM) (Macaca silenus, n = 9) and ring-tailed lemurs (RTL) (Lemur catta, n = 13), residing on St. Catherines Island, GA, was observed seasonally at quarterly intervals for 1 year. Each group had unlimited access to native flora and fauna in addition to a daily provisioned diet; LTM were observed a total of 144 hours and RTL for 208 hours. We hypothesized that foods selected by macaques would be lower in macronutrients and higher in fiber than those consumed by lemurs owing to body size differences. Samples of plant foods observed eaten (106 samples from 30 identified species) were collected, air-dried in the field, and analyzed for macronutrient, fiber, and mineral content. Leaves and seeds were important diet components for semi free-ranging LTM most of the year, comprising 51% to 93% of observed feeding time on non-provisioned food items, with ripe fruit a major food item (40% of observed feeding time) only during the August sampling period. In general, RTL consumed a more diverse diet of fruits and leaves during all seasons, with seeds and buds increasingly important during winter months (45% of observed feeding time). Based on the nutrient composition of different plant parts, weighted by the percentage contribution (feeding time assumed equivalent to mass of forage intake) of each part, no difference was seen in fiber (LTM 56.8 ± 21.9% of DM, RTL 38.4 ± 16.7% DM) or crude protein content of diets consumed (LTM 9.0 ± 2.9% DM, RTL 11.5± 6.7% DM) throughout the year. Weighted diets contained mineral concentrations meeting recommended values for non-human primates of Mg, Cu, Mn, and Zn, but were low in Na, P, and Fe, as well as Ca (LTM only). Despite wide differences in individual food items selected, overall diets were remarkably similar in macronutrient composition between these two disparate primates. Zoo Biol 18:481–494, 1999. © 1999 Wiley-Liss, Inc.

Data are presented on ring-tailed lemur feeding ecology and resource use at the Beza Mahafaly Special Reserve in southwestern Madagascar. The phenological availability of food resources was sampled monthly from 199 trees and 31 species. Results indicate that Lemur catta feeding ecology is finely tuned to the seasonal nature of specific food resources. Key species provide important food items during critical periods of the reproductive cycle. During a drought year, mortality for mothers and infants increased dramatically, providing indirect evidence that this species is highly dependent on the phenological reliability of food resources. These results indicate that in such highly seasonal habitats, the loss of any key species could have enormous impact on ring-tailed lemur demography and survival.