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Introduced species can negatively impact endemic flora and fauna. Studies have primarily utilized camera trap observations and occupancy modelling to better clarify the presence/absence and temporal overlap of endemic and exotic predators. Longitudinal data from field research sites are important as they can provide a finer understanding of predator dynamics and their effects on endemic species. One such site is the Bezà Mahafaly Special Reserve, in southern Madagascar. Protected since the 1970s, the local human population around Bezà Mahafaly Special Reserve has greatly expanded, leading to habitat disturbance in the surrounding forests and increased contact between local wildlife, people and their livestock and dogs. Here we use a combination of scat sampling, field observations of successful and attempted predations, locations of scat samples with identifiable lemur remains, and camera trap data to better assess the predator ecology at Bezà Mahafaly Special Reserve. Our results indicate that forest cats (Felis catus), are effective predators of both adult and infant lemurs and appear to be a constant mammalian predator, utilizing mammal prey more than dogs. Dogs are both predators and scavengers of lemurs. Civets focus on small prey, such as insects and rodents as well as plant material. The fosa, Cryptoprocta ferox, are also present but may not hunt in the area continuously. The killing of an adult ring-tailed lemur by two men from outside the area indicate culturally imposed taboos against lemur killing may no longer be effective given that new arrivals may not share the same local cultural restrictions. Scat sampling of exotic predators is one way to expand our understanding of exotic and endemic predator impact on lemur populations, and long-term studies with multiple assessments of predation can provide a clearer understanding of how non-endemic and endemic predators affect endangered species survival.

South Africa has six species of primates, three of which are bushbabies (family Galagidae). Very little information is available on their parasites due to the lack of longitudinal studies, although Rhipicephalus appendiculatus, Amblyomma hebraeum and Haemaphysalis elliptica were previously reported from the brown greater galago (Otolemur crassicaudatus) in South Africa. During 2014-2019, 83 O. crassicaudatus (70 live-trapped and 13 deceased animals) were checked for the presence of hard ticks, all from Limpopo Province, South Africa. Seventy-three of 83 (88 %) galagos were found to be tick-infested. Among ixodid genera, Haemaphysalis had the highest prevalence (46 % of the bushbabies), followed by Rhipicephalus (25 %) and Ixodes (18 %). In total, ten tick species were identified. Importantly, all infestations were monospecific. Ticks occurred on various body parts of bushbabies, thus no predilection site was noted. In conclusion, while previously only three ixodid species were known to infest bushbabies in South Africa, the present study showed that these animals can be parasitized by a much broader range of hard ticks., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Copyright © 2024 The Author(s). Published by Elsevier GmbH.. All rights reserved.)

Primate species exhibit considerable variation in behavior and outcomes during intergroup encounters (IGEs). Social group characteristics, like group size, and individual traits, such as sex, rank, and reproductive status, within those groups can influence both IGE engagement and outcomes. To better understand the impact of group heterogeneity on IGEs, we must examine individual strategies to elucidate individual costs and benefits of engaging in these interactions. Here, we present a descriptive study of the IGEs between two small social groups of ring-tailed lemurs (Lemur catta) on St. Catherines Island, GA, USA. We distinguish between dyadic and multiple-individual interactions between groups to compare collective and individual agonistic engagement. All encounters occurred when the East Road group (N = 7 individuals) traveled from the center of their home range to the boundary with Windmill group's (N = 5 individuals) home range, indicating that East Road might have been actively testing this boundary for access to food resources, sleeping trees, and mating opportunities. We also found notable individual variation in participation during IGEs. The daughters of the highest-ranking females were the most-engaged in dyadic intergroup and intragroup aggression, had high "win" rates during intergroup dyadic encounters, and engaged in intergroup multiple-individual interactions at high levels. These findings indicate that they might value their home range more as "potential alphas" compared to other group members. Dominant females were the most engaged in multiple-individual interactions, suggesting that they contribute heavily to collective action that might result in a gain or loss of access to resources. Finally, these two small groups might be equally matched despite the two-individual group size disparity due to individual free-riding strategies. Future research should focus on individual strategies during IGEs to characterize the complex decisions and trade-offs that influence participation., (© 2022. The Author(s), under exclusive licence to Japan Monkey Centre.)

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.

The habitats of Galago moholi are suspected to be largely fragmented, while the species is thought to be expanding further into the southernmost fringe of its range, as well as into human settlements. To date, no intraspecific molecular genetic studies have been published on G. moholi. Here we estimate the genetic diversity and connectivity of populations of G. moholi using two mitochondrial gene regions, the cytochrome C oxidase subunit I gene (COI) and the displacement loop of the control region (D-loop). Samples from five localities in northern South Africa were obtained from archived collections. The two mitochondrial DNA gene regions were amplified and sequenced to provide population summary statistics, differentiation [proportion of the total genetic variation in a population relative to the total genetic variance of all the populations (F ST ), differentiation within populations among regions (Φ ST )], genetic distance and structure. There was discernible genetic structure among the individuals, with two COI and six D-loop haplotypes belonging to two genetically different groups. There was population differentiation among regions (F ST  = 0.670; Φ ST  = 0.783; P < 0.01). However, there were low levels of differentiation among populations, as haplotypes were shared between distant populations. Adjacent populations were as divergent from each other as from distant populations. The results suggest that genetic introgression, most likely due to past migrations or recent unintentional translocations that include the animal trade, may have led to connectivity among populations.

Madagascar's ring-tailed lemurs ( Lemur catta ) are experiencing rapid population declines due to ongoing habitat loss and fragmentation, as well as increasing exploitation for bushmeat and the illegal pet trade. Despite being the focus of extensive and ongoing behavioral studies, there is comparatively little known about the genetic population structuring of the species. Here, we present the most comprehensive population genetic analysis of ring-tailed lemurs to date from across their likely remaining geographic range. We assessed levels of genetic diversity and population genetic structure using multilocus genotypes for 106 adult individuals from nine geographically representative localities. Population structure and F ST analyses revealed moderate genetic differentiation with localities being geographically partitioned into northern, southern, western and also potentially central clusters. Overall genetic diversity, in terms of allelic richness and observed heterozygosity, was high in the species (AR = 4.74, H O  = 0.811). In fact, it is the highest among all published lemur estimates to date. While these results are encouraging, ring-tailed lemurs are currently affected by ongoing habitat fragmentation and occur at lower densities in poorer quality habitats. The effects of continued isolation and fragmentation, coupled with climate-driven environmental instability, will therefore likely impede the long-term viability of the species., Competing Interests: None declared., (© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Documenting the natural diversity of eukaryotic organisms in the nonhuman primate (NHP) gut is important for understanding the evolution of the mammalian gut microbiome, its role in digestion, health and disease, and the consequences of anthropogenic change on primate biology and conservation. Despite the ecological significance of gut-associated eukaryotes, little is known about the factors that influence their assembly and diversity in mammals. In this study, we used an 18S rRNA gene fragment metabarcoding approach to assess the eukaryotic assemblage of 62 individuals representing 16 NHP species. We find that cercopithecoids, and especially the cercopithecines, have substantially higher alpha diversity than other NHP groups. Gut-associated protists and nematodes are widespread among NHPs, consistent with their ancient association with NHP hosts. However, we do not find a consistent signal of phylosymbiosis or host-species specificity. Rather, gut eukaryotes are only weakly structured by primate phylogeny with minimal signal from diet, in contrast to previous reports of NHP gut bacteria. The results of this study indicate that gut-associated eukaryotes offer different information than gut-associated bacteria and add to our understanding of the structure of the gut microbiome.

Greater bushbabies, strepsirrhine primates, that are distributed across central, eastern and southern Africa, with northern and eastern South Africa representing the species' most southerly distribution. Greater bushbabies are habitat specialists whose naturally fragmented habitats are getting even more fragmented due to anthropogenic activities. Currently, there is no population genetic data or study published on the species. The aim of our study was to investigate the genetic variation in a thick-tailed bushbaby, Otolemur crassicaudatus, population in the Soutpansberg mountain range, Limpopo Province, South Africa. Four mitochondrial regions, ranging from highly conserved to highly variable, were sequenced from 47 individuals. The sequences were aligned and genetic diversity, structure, as well as demographic analyses were performed. Low genetic diversity (π = 0.0007-0.0038 in coding regions and π = 0.0127 in non-coding region; Hd = 0.166-0.569 in coding regions and Hd = 0.584 in non-coding region) and sub-structuring ( H  = 2-3 in coding regions and H  = 4 in non-coding region) was observed with two divergent haplogroups (haplotype pairwise distance = 3-5 in coding region and 6-10 in non-coding region) being identified. This suggests the population may have experienced fixation of mitochondrial haplotypes due to limited female immigration, which is consistent with philopatric species, that alternative haplotypes are not native to this population, and that there may be male mobility from adjacent populations. This study provides the first detailed insights into the mitochondrial genetic diversity of a continental African strepsirrhine primate and demonstrates the utility of mitochondrial DNA in intraspecific genetic population analyses of these primates.

The health of 44 wild ring-tailed lemurs ( Lemur catta) at the Bezà Mahafaly Special Reserve was assessed across three age classes: <5 yr (young), 5-9 yr (adult), and ≥10 yr (old). Hematology and biochemistry tests were performed manually (leukocyte count and differential, packed cell volume, total protein) and using a point-of-care analyzer (hematocrit, hemoglobin, glucose, blood urea nitrogen, creatinine, sodium, potassium, chloride, ionized calcium, total carbon dioxide, anion gap), respectively. Urine specific gravity was measured via refractometry. Age- and sex-related differences were detected. Old lemurs had significantly lower lymphocyte count than adult and young lemurs, leading to markedly lower total leukocyte count and higher neutrophil-to-lymphocyte ratio. Decreased lymphocyte count with advanced age is consistent with immunosenescence. Young lemurs had significantly higher total protein, monocyte count, and potassium than adult and old lemurs but significantly lower ionized calcium than adult lemurs. Males had significantly higher leukocyte, neutrophil, and monocyte counts; lower percentage basophils; and higher blood urea nitrogen than females. Females had markedly higher glucose than males. Young females had the highest monocyte count and total protein, which were significantly lower in the adult and old age classes. Basophil count was stable in females across age but dropped precipitously in males in the adult and old age classes. Within adult and old age classes, males had significantly higher blood urea nitrogen and lower basophils than females. Glucose was significantly higher after α2 agonist administration. Identifying age-related hematologic and biochemical changes in apparently healthy wild ring-tailed lemurs will aid in clinical diagnosis and treatment of lemurs in human care, which is especially relevant for management of geriatric animals in zoo populations. Equally important, a better understanding of the ability of aging lemurs to tolerate environmental stressors will inform the capacity for this species to cope with ongoing and future habitat alteration.

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 Bezà 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., (© 2017 Wiley Periodicals, Inc.)

Leprosy is caused by the bacterial pathogens Mycobacterium leprae and Mycobacterium lepromatosis. Apart from humans, animals such as nine-banded armadillos in the Americas and red squirrels in the British Isles are naturally infected with M. leprae. Natural leprosy has also been reported in certain nonhuman primates, but it is not known whether these occurrences are due to incidental infections by human M. leprae strains or by M. leprae strains specific to nonhuman primates. In this study, complete M. leprae genomes from three naturally infected nonhuman primates (a chimpanzee from Sierra Leone, a sooty mangabey from West Africa, and a cynomolgus macaque from The Philippines) were sequenced. Phylogenetic analyses showed that the cynomolgus macaque M. leprae strain is most closely related to a human M. leprae strain from New Caledonia, whereas the chimpanzee and sooty mangabey M. leprae strains belong to a human M. leprae lineage commonly found in West Africa. Additionally, samples from ring-tailed lemurs from the Bezà Mahafaly Special Reserve, Madagascar, and chimpanzees from Ngogo, Kibale National Park, Uganda, were screened using quantitative PCR assays, to assess the prevalence of M. leprae in wild nonhuman primates. However, these samples did not show evidence of M. leprae infection. Overall, this study adds genomic data for nonhuman primate M. leprae strains to the existing M. leprae literature and finds that this pathogen can be transmitted from humans to nonhuman primates as well as between nonhuman primate species. While the prevalence of natural leprosy in nonhuman primates is likely low, nevertheless, future studies should continue to explore the prevalence of leprosy-causing pathogens in the wild.

Across species, diversity at the major histocompatibility complex (MHC) is critical to individual disease resistance and, hence, to population health; however, MHC diversity can be reduced in small, fragmented, or isolated populations. Given the need for comparative studies of functional genetic diversity, we investigated whether MHC diversity differs between populations which are open, that is experiencing gene flow, versus populations which are closed, that is isolated from other populations. Using the endangered ring-tailed lemur ( Lemur catta ) as a model, we compared two populations under long-term study: a relatively "open," wild population ( n  = 180) derived from Bezà Mahafaly Special Reserve, Madagascar (2003-2013) and a "closed," captive population ( n  = 121) derived from the Duke Lemur Center (DLC, 1980-2013) and from the Indianapolis and Cincinnati Zoos (2012). For all animals, we assessed MHC-DRB diversity and, across populations, we compared the number of unique MHC-DRB alleles and their distributions. Wild individuals possessed more MHC-DRB alleles than did captive individuals, and overall, the wild population had more unique MHC-DRB alleles that were more evenly distributed than did the captive population. Despite management efforts to maintain or increase genetic diversity in the DLC population, MHC diversity remained static from 1980 to 2010. Since 2010, however, captive-breeding efforts resulted in the MHC diversity of offspring increasing to a level commensurate with that found in wild individuals. Therefore, loss of genetic diversity in lemurs, owing to small founder populations or reduced gene flow, can be mitigated by managed breeding efforts. Quantifying MHC diversity within individuals and between populations is the necessary first step to identifying potential improvements to captive management and conservation plans.

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., (© 2016 The Authors. American Journal of Primatology Published by Wiley Periodicals, Inc.)

Determining the proximate causes of tooth wear remains a major focus of dental study. Here we compare the diets of three ring-tailed lemur (Lemur catta) populations and examine how different dietary components may contribute to patterns of wear-related tooth shape. Casts were made from dental impressions collected between 2003 and 2010 from lemurs in the gallery and spiny/mixed forests of the Bezá Mahafaly Special Reserve (BMSR; Parcels 1 and 2) and the spiny/mixed forests of Tsimanampesotse National Park (TNP), Madagascar. Tooth shape variables (occlusal relief and slope, angularity) were analyzed using dental topographic analysis. Focal observations and food mechanical properties (FMPs: toughness, hardness, elastic modulus) were conducted and tested, respectively, during wet and dry seasons from 2008 to 2012. We found that FMPs correlate with patterns of dental topography in these three populations. Specifically, food toughness and elastic modulus correlate with the dental variables, but hardness does not. Average food toughness and elastic modulus, but not hardness, are highest in BMSR Parcel 2, followed by BMSR Parcel 1 and TNP. Occlusal relief and slope, which serve as proxies for tooth wear, show the greatest wear in Parcel 2 and the least in TNP. Angularity is also more pronounced in TNP. Further, dental topographic patterns correspond to reliance on Tamarindus indica (tamarind) fruit. Both BMSR populations consume tamarind at high frequencies in the dry season, but the fruits are rare at TNP and only occasionally consumed. Thus, high seasonal tamarind consumption and its mechanical values help explain the low dental relief and slope among BMSR lemurs. By investigating the ecology of a single widespread species across a variety of habitats, we have been able to link specific components of diet to patterns of dental topography in this species. This provides a context for interpreting wear-related tooth shape changes more generally, illustrating that populations can develop different dental wear patterns resulting from a mix of intrinsic factors (thin enamel) and local conditions (food properties, frequency of consumption)., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The gut microbiota contributes to host health by maintaining homeostasis, increasing digestive efficiency, and facilitating the development of the immune system. The composition of the gut microbiota can change dramatically within and between individuals of a species as a result of diet, age, or habitat. Therefore, understanding the factors determining gut microbiota diversity and composition can contribute to our knowledge of host ecology as well as to conservation efforts. Here we use high-throughput sequencing to describe variation in the gut microbiota of the endangered ring-tailed lemur (Lemur catta) at the Bezà Mahafaly Special Reserve (BMSR) in southwestern Madagascar. Specifically, we measured the diversity and composition of the gut microbiota in relation to social group, age, sex, tooth wear and loss, and habitat disturbance. While we found no significant variation in the diversity of the ring-tailed lemur gut microbiota in response to any variable tested, the taxonomic composition of the gut microbiota was influenced by social group, age, and habitat disturbance. However, effect sizes were small and appear to be driven by the presence or absence of relatively low abundance taxa. These results suggest that habitat disturbance may not impact the lemur gut microbiota as strongly as it impacts the gut microbiota of other primate species, highlighting the importance of distinct host ecological and physiological factors on host-gut microbe relationships. Am. J. Primatol. 78:883-892, 2016. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Background: Across species, diversity at the Major Histocompatibility Complex (MHC) is critical to disease resistance and population health; however, use of MHC diversity to quantify the genetic health of populations has been hampered by the extreme variation found in MHC genes. Next generation sequencing (NGS) technology generates sufficient data to genotype even the most diverse species, but workflows for distinguishing artifacts from alleles are still under development. We used NGS to evaluate the MHC diversity of over 300 captive and wild ring-tailed lemurs (Lemur catta: Primates: Mammalia). We modified a published workflow to address errors that arise from deep sequencing individuals and tested for evidence of selection at the most diverse MHC genes., Results: In addition to evaluating the accuracy of 454 Titanium and Ion Torrent PGM for genotyping large populations at hypervariable genes, we suggested modifications to improve current methods of allele calling. Using these modifications, we genotyped 302 out of 319 individuals, obtaining an average sequencing depth of over 1000 reads per amplicon. We identified 55 MHC-DRB alleles, 51 of which were previously undescribed, and provide the first sequences of five additional MHC genes: DOA, DOB, DPA, DQA, and DRA. The additional five MHC genes had one or two alleles each with little sequence variation; however, the 55 MHC-DRB alleles showed a high dN/dS ratio and trans-species polymorphism, indicating a history of positive selection. Because each individual possessed 1-7 MHC-DRB alleles, we suggest that ring-tailed lemurs have four, putatively functional, MHC-DRB copies., Conclusions: In the future, accurate genotyping methods for NGS data will be critical to assessing genetic variation in non-model species. We recommend that future NGS studies increase the proportion of replicated samples, both within and across platforms, particularly for hypervariable genes like the MHC. Quantifying MHC diversity within non-model species is the first step to assessing the relationship of genetic diversity at functional loci to individual fitness and population viability. Owing to MHC-DRB diversity and copy number, ring-tailed lemurs may serve as an ideal model for estimating the interaction between genetic diversity, fitness, and environment, especially regarding endangered species.

Genetic variability among captive and wild ring-tailed lemurs (Lemur catta) was assessed using mitochondrial and nuclear DNA data. A 529 bp segment of mtDNA was sequenced and 9 microsatellite loci were genotyped for 286 ring-tailed lemurs. Samples were obtained from the well-studied L. catta population at the Bezà Mahafaly Special Reserve and from captive animals at six institutions worldwide. We found evidence of possible patrilineal contribution but the absence of matrilineal contribution from the Bezà area, and haplotypes not found in Bezà but present in Ambohimahavelona, Andringitra Massif, and other unknown locations, in the sampled captive population, indicating that the founders of the captive population originated from a wide geographic range. Total genetic variation and relatedness in captive L. catta in the six institutions were similar in extent to that of the wild population in Bezà. Based on the diverse origins of the captive population founders our results suggest the erosion of genetic diversity in the captive population. Sampled individuals from the same institution were more closely related to each other than members of a social group in the wild. Individuals housed at different institutions were less closely related than those of different social groups at Bezà, indicating lower genetic exchange between captive institutions than between social groups in a locality in the wild. Our findings underscore the usefulness of genotyping in determining the geographic origin of captive population founders, obtaining pedigree information if paternity is uncertain, and in maximizing preservation of extant genetic diversity in captivity., (© 2015 Wiley Periodicals, Inc.)

The ring-tailed lemur (Lemur catta) is a group-living strepsirrhine primate endemic to Madagascar that faces considerable predation pressure from aerial and terrestrial predators. This species engages in mobbing and vigilance behavior in response to predators, and has referential alarm vocalizations. Because L. catta is female dominant, less is known about the alarm calls of males. We tested 3 hypotheses for male antipredator vocalization behavior on L. catta at the Bezà Mahafaly Special Reserve in Madagascar: the predator confusion, group maintenance, and predation risk allocation hypotheses. We found support for 2 hypotheses. When a male L. catta made an antipredator call, other group members vocalized in response. Dominant males did not make alarm calls at higher rates than subordinate males. Predators were more abundant on the western side of Parcel 1, but an even greater number of antipredator vocalizations occurred in this area than predator abundance warranted. We show that male L. catta consistently participated in group-level antipredator vocalization usage in high-risk locations. Although female L. catta are known to hold the primary role in group defense, male L. catta are also key participants in group-wide behaviors that may confuse or drive away predators., (© 2015 S. Karger AG, Basel.)

Ring-tailed lemurs have been studied intensively in the Parcel 1 gallery forest of Bezà Mahafaly Special Reserve. Here, we report on lemur groups in a mixture of deciduous dry forest and spiny forest just 5 km to the west. Compared to Parcel 1, Parcel 2 (P2) has a lower density of Tamarindus indica, a major dietary plant species for gallery forest lemurs. Recent studies in drier habitats have called into question the association of lemur density and tamarind presence. In order to address this question, we measured forest structure and composition of plant plots between parcels and conducted lemur feeding observations. The trees and shrubs within the parcels did not differ in height or diameter at breast height, but the frequencies of plant species that were common between parcels were significantly different. Numbers of feeding observations on foods common to both parcels did not differ, but their relative rankings within parcels did. Frequencies of food plants corresponded to earlier reports of lemur population densities. However, we found that the ring-tailed lemur diet is a mixture of plants that are eaten in abundance regardless of frequency and those that are locally available. In terms of their reliance on Tamarindus, P2 animals appear intermediate between those in gallery forests and nontamarind sites., (© 2015 S. Karger AG, Basel.)

Coat and body mass status provide a potential noninvasive way to assess primate health status as well as the effects of seasonality, resource use and reproductive state. Coat and body condition were scored visually for 36 wild Lemur catta at the Bezà Mahafaly Special Reserve, Madagascar, from July 2012 to March 2013. Coat quality generally increased during the wet season when resource availability increased, in contrast to that observed during the resource-depleted dry season. Alopecia frequency increased from June to December and declined between January and March. Sex differences for coat condition were only observed in January, when males had superior coat scores. Body condition did not vary by month or sex except in February, when males were larger than females. Females that birthed infants were of lower body size than individuals who did not for November and from January to March. Our results indicate visual methods effectively detect variability in coat and body condition related to seasonality and reproductive status. Such methods present a noninvasive means for assessing the impact of seasonal resource availability, stresses of infant care and reproductive state on ring-tailed lemurs, and may be useful for assessing the impacts of these factors on general health status., (© 2015 S. Karger AG, Basel.)

Although Lemur catta persists in many habitat types in southern Madagascar, its ecology has been primarily studied within gallery forests. We compare plant food selection and properties for ring-tailed lemurs in the spiny and gallery forests over the synchronized lactation period (September to March) that includes both the dry and wet seasons. We found no significant habitat-specific differences in the type of plant part consumed per month (i.e. flower, fruit, leaf) or between the intake of soluble carbohydrates. However, the presence and use of Tamarindus indica plants appear to elevate protein and fiber intake in the gallery forest lemurs' diets. Protein is especially important for reproductive females who incur the added metabolic costs associated with lactation; however, fiber can disrupt protein digestion. Future work should continue to investigate how variations of protein and fiber affect ring-tailed lemur dietary choice and nutrient acquisition., (© 2015 S. Karger AG, Basel.)

The Ankilitelo cave site, Madagascar, contains a large collection of extant and recently extinct subfossil lemurs including the extant taxa Lemur catta and Eulemur rufifrons, which today are rarely found in sympatry. Dates for this assemblage range from 300 to 13,000 BP, though known dates for extinct primate specimens range between ∼500 and ∼600 BP. Data from Ankilitelo L. catta and E. rufifrons were compared to assess tooth wear in sympatric, related forms. Wear was scored using an ordinal scale from 0 to 5. For P4, M1 and M2, E. rufifrons displays significantly more wear than L. catta. Ankilitelo represents one of the most southerly samples of E. rufifrons, and wear data suggest that in the recent (i.e. Holocene) past, their diet near the edges of their geographic range included mechanically challenging foods. In contrast, sympatric L. catta was using foods in this transitional humid-dry forest with succulent woodlands that were not significantly impacted by recent human actions, and for which they were dentally adapted. Results also suggest that this non-gallery forest habitat may be the 'adaptive home' of L. catta, given the lack of notable tooth wear when compared to populations currently living in tamarind-dominated riverine gallery forests., (© 2015 S. Karger AG, Basel.)

The health of 36 wild, free-ranging ring-tailed lemurs (Lemur catta) at the Bezà Mahafaly Special Reserve was assessed across 2 habitats of varied human impact: a reserve riverine gallery forest, and a degraded mixed dry deciduous and Alluaudia-dominated spiny forest. While there were no statistically significant differences in leukocyte count or differential between habitats, female lemurs in the reserve gallery forest had significantly higher percentages of monocytes and eosinophils than male lemurs in the gallery forest. Lemurs from the degraded spiny habitat had significantly higher mean packed cell volume, hematocrit, hemoglobin, total protein, blood urea nitrogen, chloride, ionized calcium and urine specific gravity than lemurs from the reserve gallery forest. These findings may reflect lower hydration levels in lemurs living in degraded habitat, providing evidence that environmental degradation has identifiable impacts on the physiology and health of wild, free-ranging ring-tailed lemurs living in nearby habitats. Given the greater evidence of human impact in the mixed dry deciduous/spiny forest habitat, a pattern seen throughout southern Madagascar, biomedical markers suggestive of decreased hydration can provide empirical data to inform new conservation policies facilitating the long-term survival of this lemur community., (© 2015 S. Karger AG, Basel.)

Lemur catta has traditionally been considered a species with male-biased dispersal; however, occasional female dispersal occurs. Using molecular data, we evaluated dispersal patterns in 2 L. catta populations in southwestern Madagascar: Tsimanampesotse National Park (TNP) and Bezà Mahafaly Special Reserve (BMSR). We also investigated the genetic differentiation between the populations and dispersal partner relatedness. Results showed minor genetic differentiation between the populations (ϴ(ST) = 0.039), which may indicate gene flow historically occurring in this region, made possible by the presence of L. catta groups between the sites. Different patterns of sex-biased dispersal were found between the sites using corrected assignment indices: male-biased dispersal in TNP, and a lack of sex-biased dispersal in BMSR. Observational evidence of female dispersal in BMSR supports these results and may imply intense female resource competition in and around BMSR, because small groups of 2-3 females have been observed dispersing within BMSR and entering the reserve from outside. These dispersing groups largely consisted of mothers transferring with daughters, although we have an aunt-niece pair transferring together. Genetic data suggest that males also transfer with relatives. Our data demonstrate that dispersal partners consist of same-sexed kin for L. catta males and females, highlighting the importance of kin selection., (© 2015 S. Karger AG, Basel.)

For over 50 years, ring-tailed lemurs have been studied continuously in the wild. As one of the most long-studied primate species, the length and breadth of their study is comparable to research on Japanese macaques, baboons and chimpanzees. They are also one of the most broadly observed of all primates, with comprehensive research conducted on their behaviour, biology, ecology, genetics, palaeobiology and life history. However, over the last decade, a new generation of lemur scholars, working in conjunction with researchers who have spent decades studying this species, have greatly enhanced our knowledge of ring-tailed lemurs. In addition, research on this species has expanded beyond traditional gallery forest habitats to now include high altitude, spiny thicket, rocky outcrop and anthropogenically disturbed coastal forest populations. The focus of this special volume is to 're-imagine' the 'flagship species of Madagascar', bringing together three generations of lemur scholars., (© 2015 S. Karger AG, Basel.)

Ring-tailed lemurs (Lemur catta) at the Bezà 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., (© 2014 Wiley Periodicals, Inc.)

A thorough knowledge of biological variation in extant primates is imperative for interpreting variation, and for delineating species in primate biology and paleobiology. This is especially the case given the recent, rapid taxonomic expansion in many primate groups, notably among small-bodied nocturnal forms. Here we present data on dental, cranial, and pelage variation in a single-locality museum sample of mouse lemurs from Amboasary, Madagascar. To interpret these data, we include comparative information from other museum samples, and from a newly collected mouse lemur skeletal sample from the Beza Mahafaly Special Reserve (BMSR), Madagascar. We scored forty dental traits (n = 126) and three pelage variants (n = 19), and collected 21 cranial/dental measures. Most dental traits exhibit variable frequencies, with some only rarely present. Individual dental variants include misshapen and supernumerary teeth. All Amboasary pelage specimens display a "reversed V" on the cap, and a distinct dorsal median stripe on the back. All but two displayed the dominant gray-brown pelage coloration typical of Microcebus griseorufus. Cranial and dental metric variability are each quite low, and craniometric variation does not illustrate heteroscedasticity. To assess whether this sample represents a single species, we compared dental and pelage variation to a documented, single-species M. griseorufus sample from BMSR. As at Amboasary, BMSR mouse lemurs display limited odontometric variation and wide variation in non-metric dental traits. In contrast, BMSR mouse lemurs display diverse pelage, despite reported genetic homogeneity. Ranges of dental and pelage variation at BMSR and Amboasary overlap. Thus, we conclude that the Amboasary mouse lemurs represent a single species - most likely (in the absence of genetic data to the contrary) M. griseorufus, and we reject their previous allocation to Microcebus murinus. Patterns of variation in the Amboasary sample provide a comparative template for recognizing the degree of variation manifested in a single primate population, and by implication, they provide minimum values for this species' intraspecific variation. Finally, discordance between different biological systems in our mouse lemur samples illustrates the need to examine multiple systems when conducting taxonomic analyses among living or fossil primates., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Teeth are central for the study of ecology, as teeth are at the direct interface between an organism and its environment. Recent years have witnessed a rapid growth in the use of teeth to understand a broad range of topics in living and fossil primate biology. This in part reflects new techniques for assessing ways in which teeth respond to, and interact with, an organism's environment. Long-term studies of wild primate populations that integrate dental analyses have also provided a new context for understanding primate interactions with their environments. These new techniques and long-term field studies have allowed the development of a new perspective-dental ecology. We define dental ecology as the broad study of how teeth respond to, or interact with, the environment. This includes identifying patterns of dental pathology and tooth use-wear, as they reflect feeding ecology, behavior, and habitat variation, including areas impacted by anthropogenic disturbance, and how dental development can reflect environmental change and/or stress. The dental ecology approach, built on collaboration between dental experts and ecologists, holds the potential to provide an important theoretical and practical framework for inferring ecology and behavior of fossil forms, for assessing environmental change in living populations, and for understanding ways in which habitat impacts primate growth and development. This symposium issue brings together experts on dental morphology, growth and development, tooth wear and health, primate ecology, and paleontology, to explore the broad application of dental ecology to questions of how living and fossil primates interact with their environments., (Copyright © 2012 Wiley Periodicals, Inc.)

Knowledge of dietary mechanical properties can be informative about physical consequences to consumers during ingestion and mastication. In this article, we examine how Tamarindus indica fruits can affect dental morphology in a population of ring-tailed lemurs (Lemur catta) at Beza Mahafaly special reserve in southwestern Madagascar. Ring-tailed lemurs in tamarind dominated gallery forests exhibit extreme wear and tooth loss on their postcanine dentition that has been related to processing T. indica fruits. We measured and compared mechanical properties of individual food parts in the diet of ring-tailed lemurs in different seasons in 1999-2000, 2008, and 2010. Fracture toughness, hardness, and modulus of foods were measured with a portable mechanical tester. The ripe fruits of T. indica are indeed the toughest and hardest foods ingested by the lemurs. In addition, they are among the largest foods consumed, require high numbers of ingestive bites to process, and are the most frequently eaten by volume. During controlled cutting tests of the ripe fruit shell, multiple runaway side cracks form alongside the cut. Similarly, the lemurs repeatedly bite the ripe shell during feeding and thereby introduce multiple cracks that eventually fragment the shell. Studies of enamel microstructure (e.g., Lucas et al.: BioEssays 30 (2008) 374-385; Campbell et al., 2011) advance the idea that the thin enamel of ring-tailed lemur teeth is susceptible to substantial micro-cracking that rapidly erodes the teeth. We conclude that micro-cracking from repeated loads, in combination with the mechanical and physical properties of the fruit, is primarily responsible for the observed dental damage., (Copyright © 2012 Wiley Periodicals, Inc.)

The common morphological metrics of size, shape, and enamel thickness of teeth are believed to reflect the functional requirements of a primate's diet. However, the mechanical and material properties of enamel also contribute to tooth function, yet are rarely studied. Substantial wear and tooth loss previously documented in Lemur catta at the Beza Mahafaly Special Reserve suggests that their dental morphology, structure, and possibly their enamel are not adapted for their current fallback food (the mechanically challenging tamarind fruit). In this study, we investigate the nanomechanical properties, mineralization, and microstructure of the enamel of three sympatric lemur species to provide insight into their dietary functional adaptations. Mechanical properties measured by nanoindentation were compared to measurements of mineral content, prism orientation, prism size, and enamel thickness using electron microscopy. Mechanical properties of all species were similar near the enamel dentin junction and variations correlated with changes in microstructure (e.g., prism size) and mineral content. Severe wear and microcracking within L. catta's enamel were associated with up to a 43% reduction in nanomechanical properties in regions of cracking versus intact enamel. The mechanical and material properties of L. catta's enamel are similar to those of sympatric folivores and suggest that they are not uniquely mechanically adapted to consume the physically challenging tamarind fruit. An understanding of the material and mechanical properties of enamel is required to fully elucidate the functional and ecological adaptations of primate teeth., (Copyright © 2012 Wiley Periodicals, Inc.)

Teeth have long been used as indicators of primate ecology. Early work focused on the links between dental morphology, diet, and behavior, with more recent years emphasizing dental wear, microstructure, development, and biogeochemistry, to understand primate ecology. Our study of Lemur catta at the Beza Mahafaly Special Reserve, Madagascar, has revealed an unusual pattern of severe tooth wear and frequent tooth loss, primarily the result of consuming a fallback food for which these primates are not dentally adapted. Interpreting these data was only possible by combining our areas of expertise (dental anatomy [FC] and primate ecology [MS]). By integrating theoretical, methodological, and applied aspects of both areas of research, we adopted the term "dental ecology"-defined as the broad study of how teeth respond to the environment. Specifically, we view dental ecology as an interpretive framework using teeth as a vehicle for understanding an organism's ecology, which builds upon earlier work, but creates a new synthesis of anatomy and ecology that is only possible with detailed knowledge of living primates. This framework includes (1) identifying patterns of dental pathology and tooth use-wear, within the context of feeding ecology, behavior, habitat variation, and anthropogenic change, (2) assessing ways in which dental development and biogeochemical signals can reflect habitat, environmental change and/or stress, and (3) how dental microstructure and macro-morphology are adapted to, and reflect feeding ecology. Here we define dental ecology, provide a short summary of the development of this perspective, and place our new work into this context., (Copyright © 2012 Wiley Periodicals, Inc.)

During mastication, foods are reduced into particles suitable for swallowing and digestion. Smaller particles possess a greater surface area per unit of volume on which digestive enzymes and bacteria may work than relatively larger particles, and are thus more readily digested. As dental morphology facilitates the breakdown of diets with specific mechanical properties, extensive dental wear and/or tooth loss may impede an individual's ability to break down and exploit foods. We present data demonstrating a relationship between dental impairment and particle size in 43 fecal samples from 33 ring-tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar. All fecal samples were sifted through three sieves of decreasing size (11.2 mm, 4.75 mm, and 1.0 mm). The resulting fraction in each sieve was then weighed and assessed in relation to individual dental impairment status. With increasing wear, the percentage of each sample within the 1.0 mm sieve decreases, whereas that in the 11.2 mm sieve increases with increasing postcanine wear, although these effects are not present when limited to individuals without tooth loss. Individuals with tooth loss also demonstrate larger proportions of fecal material 1.0-4.75 mm in size. Dental impairment results in larger food particles and potentially less efficient utilization of foods. When fecal material was examined by leaf vs. fruit content, individuals with tooth loss demonstrated reduced proportions of fruit in the 1.0 mm and 11.2 mm sieves. These data suggest individuals with tooth loss consume less fruit than those without loss, potentially reflecting a reduced ability to process tamarind fruit, a key fallback resource at BMSR., (Copyright © 2012 Wiley Periodicals, Inc.)

In light of historical and recent anthropogenic influences on Malagasy primate populations, in this study ring-tailed lemur (Lemur catta) samples from two sites in southwestern Madagascar, Beza Mahafaly Special Reserve (BMSR) and Tsimanampetsotsa National Park (TNP), were evaluated for the genetic signature of a population bottleneck. A total of 45 individuals (20 from BMSR and 25 from TNP) were genotyped at seven microsatellite loci. Three methods were used to evaluate these populations for evidence of a historical bottleneck: M-ratio, mode-shift, and heterozygosity excess tests. Three mutation models were used for heterozygosity excess tests: the stepwise mutation model (SMM), two-phase model (TPM), and infinite allele model (IAM). M-ratio estimations indicated a potential bottleneck in both populations under some conditions. Although mode-shift tests did not strongly indicate a population bottleneck in the recent historical past when samples from all individuals were included, a female-only analysis indicated a potential bottleneck in TNP. Heterozygosity excess was indicated under two of the three mutation models (IAM and TPM), with TNP showing stronger evidence of heterozygosity excess than BMSR. Taken together, these results suggest that a bottleneck may have occurred among L. catta in southwestern Madagascar in the recent past. Given knowledge of how current major stochastic climatic events and human-induced change can negatively impact extant lemur populations, it is reasonable that comparable events in the historical past could have caused a population bottleneck. This evaluation additionally functions to highlight the continuing environmental and anthropogenic challenges faced by lemurs in southwestern Madagascar., (Copyright © 2011 Wiley Periodicals, Inc.)

Wild ring-tailed lemurs (Lemur catta) can be anesthetized with Telazol via blow dart, but improved techniques are needed so that each lemur is reliably induced with a single dart. Medetomidine-butorphanol (MB) is a good supplemental protocol to be administered once the lemurs are captured, but other protocols may provide longer periods of sedation and immobility. One possible way of increasing the efficacy of each dart is to increase the time it is retained in the leg. In this investigation, a "double-sleeve" technique was used to try to increase the time of dart retention. This technique used a standard silicone sleeve on the needle, along with a second sleeve at the needle hub. Induction values were compared between lemurs darted with double-sleeve needles and those induced with needles that each had a single silicone sleeve. Once the lemurs were induced, supplementation with MB (0.04 mg/kg and 0.2 mg/kg) was compared with supplementation with ketamine-medetomidine (KM) (10 mg/ kg and 0.04 mg/kg). Twenty-three lemurs were darted with Telazol by using single-sleeve needles, and 24 were darted with double-sleeve needles. The number of darts per lemur and number of escapes were not different between animals darted with a single sleeve compared with a double-sleeve; thus, there were no significant improvements in induction success with the double-sleeve technique. Adequate sedation and muscle relaxation were achieved with both MB and KM; however, lemurs that received MB were more relaxed and needed fewer additional supplements that those that received KM. Single-sleeve dart needles are recommended for Telazol induction of ring-tailed lemurs via blow dart and MB is preferable to KM for supplemental sedation and muscle relaxation.

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 values < 90%. 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.

Tooth wear is generally an age-related phenomenon, often assumed to occur at similar rates within populations of primates and other mammals, and has been suggested as a correlate of reduced offspring survival among wild lemurs. Few long-term wild studies have combined detailed study of primate behavior and ecology with dental analyses. Here, we present data on dental wear and tooth loss in older (>10 years old) wild and captive ring-tailed lemurs (Lemur catta). Among older ring-tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar (n=6), the percentage of severe dental wear and tooth loss ranges from 6 to 50%. Among these six individuals, the oldest (19 years old) exhibits the second lowest frequency of tooth loss (14%). The majority of captive lemurs at the Indianapolis Zoo (n=7) are older than the oldest BMSR lemur, yet display significantly less overall tooth wear for 19 of 36 tooth positions, with only two individuals exhibiting antemortem tooth loss. Among the captive lemurs, only one lemur (a nearly 29 year old male) has lost more than one tooth. This individual is only missing anterior teeth, in contrast to lemurs at BMSR, where the majority of lost teeth are postcanine teeth associated with processing specific fallback foods. Postcanine teeth also show significantly more overall wear at BMSR than in the captive sample. At BMSR, degree of severe wear and tooth loss varies in same aged, older individuals, likely reflecting differences in microhabitat, and thus the availability and use of different foods. This pattern becomes apparent before "old age," as seen in individuals as young as 7 years. Among the four "older" female lemurs at BMSR, severe wear and/or tooth loss do not predict offspring survival., (© 2010 Wiley-Liss, Inc.)

Like most of Madagascar's endemic primates, ring-tailed lemurs (Lemur catta) face a number of threats to their survival. Although habitat loss is of greatest concern, other anthropogenic factors including environmental contamination may also affect lemur health and survival. In this study, we examined ring-tailed lemurs from the Beza Mahafaly Special Reserve (BMSR), southern Madagascar for exposure to organochlorine (OC) pesticides and metals and examined differences in contaminant concentrations between sexes and among age groups, troops, and habitats. A total of 14 pesticides and 13 metals was detected in lemur blood (24 individuals) and hair (65 individuals) samples, respectively. p,p'-DDT, heptachlor, aldrin, heptachlor epoxide, endrin aldehyde, and endrin were among the most prevalent pesticides detected. Surprisingly, the persistent metabolite of p,p'-DDT, p,p'-DDE, was not detected. The most commonly detected metals were aluminum, zinc, boron, phosphorus, silicon, and copper, whereas metals considered more hazardous to wildlife (e.g. arsenic, cadmium, lead, selenium, vanadium) were not found above detection limits. Overall, concentrations of OC pesticides and metals were low and similar to those considered to be background concentrations in other studies examining the ecotoxicology of wild mammals. Few inter-sex, -age, -troop, and -habitat differences in contaminant concentrations were observed, suggesting a uniform distribution of contaminants within the reserve. Several statistically significant relationships between lemur body size and contaminant concentrations were observed, but owing to the lack of supportive data regarding contaminant exposure in wild primates, the biological significance of these findings remains uncertain. Results of this study document exposure of ring-tailed lemurs at BMSR to multiple OC pesticides and metals and provide essential baseline data for future health and toxicological evaluations of lemurs and other wild primates, especially those in regions with expanding agricultural and mining operations.

Fallback foods are often viewed as central in shaping primate morphology, and influencing adaptive shifts in hominin and other primate evolution. Here we argue that fruit of the tamarind tree (Tamarindus indica) qualifies as a fallback food of ring-tailed lemurs (Lemur catta) at the Beza Mahafaly Special Reserve (BMSR), Madagascar. Contrary to predictions that fallback foods may select for dental and masticatory morphologies adapted to processing these foods, consumption of tamarind fruit by these lemurs leaves a distinct pattern of dental pathology among ring-tailed lemurs at BMSR. Specifically, the physical and mechanical properties of tamarind fruit likely result in a high frequency of severe tooth wear, and subsequent antemortem tooth loss, in this lemur population. This pattern of dental pathology is amplified among lemurs living in disturbed areas at Beza Mahafaly, resulting from a disproportionate emphasis on challenging tamarind fruit, due to few other fruits being available. This is in part caused by a reduction in ground cover and other plants due to livestock grazing. As such, tamarind trees remain one of the few food resources in many areas. Dental pathologies are also associated with the use of a nonendemic leaf resource Argemone mexicana, an important food during the latter part of the dry season when overall food availability is reduced. Such dental pathologies at Beza Mahafaly, resulting from the use or overemphasis of fallback foods for which they are not biologically adapted, indicate that anthropogenic factors must be considered when examining fallback foods.

Severe dental wear and tooth loss is often assumed to impede the processing, breakdown, and energetic conversion of food items, thereby negatively impacting individual health, reproduction, and survival. Ring-tailed lemurs at the Beza Mahafaly Special Reserve demonstrate exceptionally high frequencies of severe dental wear and antemortem tooth loss, yet often survive multiple years with these impairments. To test the hypothesis that these lemurs mitigate tooth loss through behavioral adjustments, we collected 191 h of observational data from 16 focal subjects, eight without tooth loss and eight with between 3% and 44% loss. These data indicate dentally-impaired ring-tailed lemurs show compensatory behaviors consistent with the demands of living in a social group. During early afternoon (12:00-14:30 h) individuals with loss showed trends towards higher frequencies of foraging and grooming, while individuals without loss rested significantly more often. Individuals with >10% loss (n = 7) showed higher frequencies of feeding, foraging, and grooming, and lower frequencies of resting during this period than individuals with <10% loss (n = 9). Individuals with tooth loss maintained relatively higher levels of feeding and foraging throughout the day. These individuals licked tamarind fruit at higher frequencies, likely spending more time softening it before ingestion. These individuals did not demonstrate longer feeding bouts overall, although bouts involving tamarinds were significantly longer. Individuals with marked toothcomb wear engaged in higher rates of certain types of allogrooming, demonstrating that social behaviors are used to compensate for reduced grooming efficiency. These data have implications for interpreting behavioral responses to dental impairment in the fossil record., (Copyright 2009 Wiley-Liss, Inc.)

Chemical deterioration of teeth is common among modern humans, and has been suggested for some extinct primates. Dental erosion caused by acidic foods may also obscure microwear signals of mechanical food properties. Ring-tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar, display frequent severe tooth wear and subsequent tooth loss. In contrast, sympatric Verreaux's sifaka display far less tooth wear and infrequent tooth loss, despite both species regularly consuming acidic tamarind fruit. We investigated the potential impact of dietary acidity on tooth wear, collecting data on salivary pH from both species, as well as salivary pH from ring-tailed lemurs at Tsimanampesotse National Park, Madagascar. We also collected salivary pH data from ring-tailed lemurs at the Indianapolis Zoo, none of which had eaten for at least 12 hr before data collection. Mean salivary pH for the BMSR ring-tailed lemurs (8.098, n=41, SD=0.550) was significantly more alkaline than Verreaux's sifaka (7.481, n=26, SD=0.458). The mean salivary pH of BMSR (8.098) and Tsimanampesotse (8.080, n=25, SD=0.746) ring-tailed lemurs did not differ significantly. Salivary pH for the Indianapolis Zoo sample (8.125, n=16, SD=0.289) did not differ significantly from either the BMSR or Tsimanampesotse ring-tailed lemurs, but was significantly more alkaline than the BMSR Verreaux's sifaka sample. Regardless of the time between feeding and collection of pH data (from several minutes to nearly 1 hr), salivary pH for each wild lemur was above the "critical" pH of 5.5, below which enamel demineralization occurs. Thus, the high pH of lemur saliva suggests a strong buffering capacity, indicating the impact of acidic foods on dental wear is short-lived, likely having a limited effect. However, tannins in tamarind fruit may increase friction between teeth, thereby increasing attrition and wear in lemurs. These data also suggest that salivary pH varies between lemur species, corresponding to broad dietary categories.

While understanding somatic variability among wild primates can provide insight into natural patterns of developmental plasticity, published data for living populations are rare. Here we provide such information for two distinct wild populations of Lemur catta. Variants observed include microtia, athelia, and female virilization. Dental variants observed include individuals with supernumerary teeth, rotated teeth, maxillary incisor agenesis, and severe malocclusion. There was a sex bias in incisor agenesis, with 5 of 7 examples (71%) found in males. The frequency of dental variants in our sample is lower than that seen in many other lemuriformes, as well as other primates. This may be a product of their less derived dental formula and/or their relatively fast dental development. Amassing such data is a critical first step to assess if wild primate populations are exhibiting normal variability or are being affected by potential inbreeding and/or environmental effects., ((c) 2007 S. Karger AG, Basel)

Stable carbon and nitrogen isotope compositions were analyzed from hair samples of 30 sympatric ring-tailed lemurs (Lemur catta) inhabiting the Beza Mahafaly Special Reserve, Madagascar. All lemurs were known individuals involved in a longitudinal study, which allowed us to explore the degree to which group membership, sex, health status, and migration influenced their stable isotope compositions. The differences in delta(13)C and delta(15)N values between groups were small (<1.5 per thousand) but highly significant. In fact, each group was tightly clustered, and discriminant function analysis of the stable isotope data assigned individuals to the group in which they were originally collared with over 90% accuracy. In general, the differences between groups reflected the degree to which they utilized forested versus open habitats. As open habitats at Beza Mahafaly often correspond to areas of anthropogenic disturbance, these data suggest that isotopic data can be useful for addressing questions of lemur conservation. There were few sex differences, but significant differences did occur between individuals of normal and suboptimal health, with those in poor health (especially those in the worst condition) being enriched in (15)N and to a lesser degree (13)C compared with healthy individuals. Moreover, lemurs that had emigrated between 2003 and 2004 had different delta(13)C and delta(15)N compositions than their original groups., ((c) 2007 Wiley-Liss, Inc.)

Coprophagy occurs in a number of animal species, including nonhuman primates. During the 2003-2004 dry seasons at the Beza Mahafaly Special Reserve, Madagascar, we observed wild ring-tailed lemurs (Lemur catta) consuming dried fecal matter from three different species. Ring-tailed lemurs consumed human feces on 12 occasions, cattle feces twice, and feral dog feces once. Coprophagy in this population may be a behavioral adaptation that provides animals access to energy and nutrients and may be an important nutritional source for older, and/or dentally impaired individuals during the dry season., ((c) 2007 Wiley-Liss, Inc.)

Complete physical examinations and biomedical sample collection were performed on 70 free-ranging ring-tailed lemurs (Lemur catta) from three different habitats in the Beza Mahfaly Special Reserve (BMSR), in southern Madagascar, to assess the impact of humans and habitat on lemur health. Lemurs were chemically immobilized with ketamine and diazepam administered via blow darts for concurrent biomedical, morphometric, and behavioral studies. Subsets of the animals had blood analyzed for hematology, serum chemistry, micronutrients, fat-soluble vitamins (vitamins A, D, and E), measures of iron metabolism, and polymerase chain reaction assays (PCR) for Toxoplasma gondii, Hemoplasma spp., Bartonella spp., Ehrlichia spp., Anaplasma phagocytophilum, and Neorickettsia risticii. Results were compared on the basis of gender and the habitats at the study site: reserve (intact gallery forest), degraded (human inhabited and altered), and marginal (dry didieracea forest with heavy grazing and tree cutting). Levels of vitamin D, triglycerides, and cholesterol, and measures of iron metabolism for BMSR lemurs were greater than those previously reported for a free-ranging lemur population (Tsimanampetsotsa Strict Nature Reserve, Madagascar) with less access to foods of anthropogenic origin. BMSR ring-tailed lemurs from a habitat with less water (marginal) had higher sodium (P = 0.051), chloride (P = 0.045), osmolality (P = 0.010), and amylase (P = 0.05) levels than lemurs from other BMSR habitats, suggesting that these lemurs were less hydrated. Vitamin D levels of male lemurs were higher (P = 0.011) than those of females at BMSR, possibly because of differences in sunning behavior or differential selection of food items. The biological significance is uncertain for other parameters with statistically significant differences. All samples tested (n = 20) were negative for the pathogens tested using PCR assays. Continued concurrent biomedical and ecological research is needed at BMSR to confirm these results and determine their association with population mortality and fecundity rates.

The ring-tailed lemurs at Beza Mahafaly Special Reserve, Madagascar, exhibit a high frequency of severe wear and antemortem tooth loss. As part of a long-term study, we collected dental data on 83 living adult ring-tailed lemurs during 2003 and 2004. Among these individuals, 192 teeth were scored as absent. The most frequently missing tooth position is M1 (24%). As M1 is the first tooth to erupt, its high frequency of absence (primarily a result of wear) is not remarkable. However, the remaining pattern of tooth loss does not correlate with the sequence of eruption. We suggest that this pattern is a function of 1) feeding ecology, as hard, tough tamarind fruit is a key fallback food of ring-tailed lemurs living in gallery forests; 2) food processing, as tamarind fruit is primarily processed in the P3-M1 region of the mouth; and 3) tooth structure, as ring-tailed lemurs possess thin dental enamel. The incongruity between thin enamel and use of a hard, tough fallback food suggests that ring-tailed lemurs living in riverine gallery forests may rely on resources not used in the past. When comparing dental health in the same individuals (n=50) between 2003 and 2004, we found that individual tooth loss can show a rapid increase over the span of one year, increasing by as much as 20%. Despite this rapid loss, individuals are able to survive, sometimes benefiting from unintentional assistance from conspecifics, from which partially processed tamarind fruit is obtained. Although less frequent in this population, these longitudinal data also illustrate that ring-tailed lemurs lose teeth due to damage and disease, similar to other nonhuman primates. The relationship between tooth loss, feeding ecology, dental structure, and individual life history in this population has implications for interpreting behavior based on tooth loss in the hominid fossil record.