Seasons of death: patterns of predation on wild lemurs and other fauna by endemic and introduced predators.

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.

Hard ticks (Acari: Ixodidae) parasitizing bushbabies (Mammalia: Galagidae) in a biodiversity hotspot of northern South Africa.

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.

Characterizing group and individual engagement in intergroup encounters between small groups of ring-tailed lemurs (Lemur catta) on St. Catherines Island, USA.

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.

Population and genetic structure of a male-dispersing strepsirrhine, Galago moholi (Primates, Galagidae), from northern South Africa, inferred from mitochondrial DNA.

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 (FST), differentiation within populations among regions (ΦST)], 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 (FST = 0.670; ΦST = 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.

Genetic population structure of endangered ring-tailed lemurs (Lemur catta) from nine sites in southern Madagascar.

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.

Population genetic structure of the thick-tailed bushbaby (Otolemur crassicaudatus) from the Soutpansberg Mountain range, Northern South Africa, based on four mitochondrial DNA regions.

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.

Biodiversity of protists and nematodes in the wild nonhuman primate gut.

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.

AGE-RELATED CHANGES IN HEMATOLOGY AND BLOOD BIOCHEMISTRY VALUES IN ENDANGERED, WILD RING-TAILED LEMURS ( LEMUR CATTA) AT THE BEZÀ MAHAFALY SPECIAL RESERVE, MADAGASCAR.

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.

Mycobacterium leprae genomes from naturally infected nonhuman primates.

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.

The effect of extreme weather events on hair cortisol and body weight in a wild ring-tailed lemur population (Lemur catta) in southwestern Madagascar.

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.

Genetic wealth, population health: Major histocompatibility complex variation in captive and wild ring-tailed lemurs (Lemur catta).

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.

Paternity in wild ring-tailed lemurs (Lemur catta): Implications for male mating strategies.

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.

Host age, social group, and habitat type influence the gut microbiota of wild ring-tailed lemurs (Lemur catta).

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.

Next-generation genotyping of hypervariable loci in many individuals of a non-model species: technical and theoretical implications.

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.

Mechanical food properties and dental topography differentiate three populations of Lemur catta in southwest Madagascar.

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).

Comparison of the genetic variation of captive ring-tailed lemurs with a wild population in Madagascar.

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.

Ring-tailed lemurs: a species re-imagined.

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.

Seasonal feeding ecology of ring-tailed lemurs: a comparison of spiny and gallery forest habitats.

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.

Beyond the Gallery Forest: Contrasting Habitat and Diet in Lemur catta Troops at Bezà Mahafaly Special Reserve.

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.

Examining visual measures of coat and body condition in wild ring-tailed lemurs at the Bezà Mahafaly Special Reserve, Madagascar.

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.