Consistency in Verreaux's sifaka home range and core area size despite seasonal variation in resource availability as assessed by Enhanced Vegetation Index (EVI).

Primates are adept at dealing with fluctuating availability of resources and display a range of responses to minimize the effects of food scarcity. An important component of primate conservation is to understand how primates adapt their foraging and ranging patterns in response to fluctuating food resources. Animals optimize resource acquisition within the home range through the selection of resource-bearing patches and choose between contrasting foraging strategies (resource-maximizing vs. area-minimizing). Our study aimed to characterize the foraging strategy of a folivorous primate, Verreaux's sifaka (Propithecus verreauxi), by evaluating whether group home range size varied between peak and lean leaf seasons within a seasonally dry tropical forest in Madagascar. We hypothesized that Verreaux's sifaka used the resource maximization strategy to select high-value resource patches so that during periods of resource depression, the home range area did not significantly change in size. We characterized resource availability (i.e., primary productivity) by season at Kirindy Mitea National Park using remotely-sensed Enhanced Vegetation Index data. We calculated group home ranges using 10 years of focal animal sampling data collected on eight groups using both 95% and 50% kernel density estimation. We used area accumulation curves to ensure each group had an adequate number of locations to reach seasonal home range asymptotes. Neither 95% home ranges nor 50% core areas differed across peak and lean leaf resource seasons, supporting the hypothesis that Verreaux's sifaka use a resource maximization strategy. With a better understanding of animal space use strategies, managers can model anticipated changes under environmental and/or anthropogenic resource depression scenarios. These findings demonstrate the value of long-term data for characterizing and understanding foraging and ranging patterns. We also illustrate the benefits of using satellite data for characterizing food resources for folivorous primates.

Evolutionary Patterns of Intersexual Power.

Dominance and leverage are both possible causes of social inequality. If sexual dimorphism influences patterns of intersexual dominance, we predicted that highly dimorphic species are constrained to exhibit male-biased power (MP), but species with low sexual dimorphism are free to demonstrate a broader range of intersexual power relationships. If market effects influence intersexual leverage, we predicted that females have more power when group composition is more male-biased and estrus is asynchronous. We analyzed data on intersexual power, sexual dimorphism, expected estrous overlap, and sex ratio for 79 extant primate species using phylogenetic logistic regression and ancestral state reconstructions. Although MP is more common, every major primate clade includes non-MP species. MP was associated with greater body mass and canine length dimorphism and with female-biased sex-ratios. Low estrous overlap was associated with non-MP. Although MP was reconstructed as likely ancestral for anthropoids, the last common ancestor of this clade probably did not exhibit high sexual dimorphism. The last common ancestor of catarrhines was probably highly dimorphic, potentially constraining intersexual power relationships. Non-MP probably evolved multiple times in primates and may be less common because multiple traits are linked to MP while fewer traits are associated with female-biased power or equality.

A Nuanced Examination of Primate Capture and Consumption and Human Socio-Economic Well-Being in Kirindy Mitea National Park, Madagascar.

The futures of human and nonhuman primates are closely tied in protected areas. Understanding this interconnectedness is especially urgent in Madagascar, one of the world's most impoverished biodiversity hotspots. Yet, no study has evaluated the relationship between poverty and lemur hunting and consumption using a composite poverty metric that includes health, education, and living standards. To address this gap, and to inform primate conservation practice and policy, we administered annual surveys to 81 households over six consecutive months (September 2018 to March 2019) in a village on the border of Kirindy Mitea National Park, Madagascar. We observed extreme deprivation scores across multiple dimensions of poverty and identified ninety-five percent of households as 'impoverished'. Of these, three-quarters (77%) of households were identified as being in 'severe poverty'. One-fifth (19%) of all households hunted lemurs and half (49%) of households consumed lemurs. While poverty eradication is an urgent need in communities around Kirindy Mitea National Park, our findings show no relationship between poverty and lemur hunting and consumption, perhaps due to the lack of variance in poverty. Our results highlight the need to investigate other contributory factors to lemur hunting and consumption locally. Because food insecurity is a known driver of lemur hunting and consumption among the study community, and because domestic meats can be preferred over protected species, we recommend testing the efficacy of livestock interventions near Kirindy Mitea National Park.

What are oxytocin assays measuring? Epitope mapping, metabolites, and comparisons of wildtype & knockout mouse urine.

Oxytocin has become a popular analyte in behavioral endocrinology in recent years, due in part to its roles in social behavior, stress physiology, and cognition. Urine samples have the advantage of being non-invasive and minimally disruptive to collect, allowing for oxytocin measurements even in some wild populations. However, methods for urinary oxytocin immunoassay have not been sufficiently optimized and rigorously assessed for their potential limitations. Using samples from oxytocin knockout (KO) and wildtype (WT) mice, we find evidence of considerable interference in unextracted urine samples, with similar distributions of measured oxytocin in both genotypes. Importantly, although this interference can be reduced by a reversed-phase solid-phase extraction (SPE), this common approach is not sufficient for eliminating false-positive signal on three immunoassay kits. To better understand the source of the observed interference, we conducted epitope mapping of the Arbor Assays antibody and assessed its cross-reactivity with known, biologically active fragments of oxytocin. We found considerable cross-reactivity (0.5-52% by-molarity) for three fragments of oxytocin that share the core epitope, with more cross-reactivity for longer fragments. Given the presence of some cross-reactivity for even the tripeptide MIF-1, it is likely that many small protein metabolites might be sufficiently similar to the epitope that at high concentrations they interfere with immunoassays. We present a new mixed-mode cation-exchange SPE method that minimizes interference-with knockout samples measuring below the assay's limit of detection-while effectively retaining oxytocin from the urine of wildtype mice. This method demonstrates good parallelism and spike recovery across multiple species (mice, dogs, sifakas, humans). Our results suggest that immunoassays of urine samples may be particularly susceptible to interference, even when using common extraction protocols, but that this interference can be successfully managed using a novel mixed-mode cation exchange extraction. These findings imply that previous conclusions based on urinary oxytocin measurements-especially those involving unextracted samples-may need to be reassessed.

Large Comparative Analyses of Primate Body Site Microbiomes Indicate that the Oral Microbiome Is Unique among All Body Sites and Conserved among Nonhuman Primates.

The study of the mammalian microbiome serves as a critical tool for understanding host-microbial diversity and coevolution and the impact of bacterial communities on host health. While studies of specific microbial systems (e.g., in the human gut) have rapidly increased, large knowledge gaps remain, hindering our understanding of the determinants and levels of variation in microbiomes across multiple body sites and host species. Here, we compare microbiome community compositions from eight distinct body sites among 17 phylogenetically diverse species of nonhuman primates (NHPs), representing the largest comparative study of microbial diversity across primate host species and body sites. Analysis of 898 samples predominantly acquired in the wild demonstrated that oral microbiomes were unique in their clustering, with distinctive divergence from all other body site microbiomes. In contrast, all other body site microbiomes clustered principally by host species and differentiated by body site within host species. These results highlight two key findings: (i) the oral microbiome is unique compared to all other body site microbiomes and conserved among diverse nonhuman primates, despite their considerable dietary and phylogenetic differences, and (ii) assessments of the determinants of host-microbial diversity are relative to the level of the comparison (i.e., intra-/inter-body site, -host species, and -individual), emphasizing the need for broader comparative microbial analyses across diverse hosts to further elucidate host-microbial dynamics, evolutionary and biological patterns of variation, and implications for human-microbial coevolution. IMPORTANCE The microbiome is critical to host health and disease, but much remains unknown about the determinants, levels, and evolution of host-microbial diversity. The relationship between hosts and their associated microbes is complex. Most studies to date have focused on the gut microbiome; however, large gaps remain in our understanding of host-microbial diversity, coevolution, and levels of variation in microbiomes across multiple body sites and host species. To better understand the patterns of variation and evolutionary context of host-microbial communities, we conducted one of the largest comparative studies to date, which indicated that the oral microbiome was distinct from the microbiomes of all other body sites and convergent across host species, suggesting conserved niche specialization within the Primates order. We also show the importance of host species differences in shaping the microbiome within specific body sites. This large, comparative study contributes valuable information on key patterns of variation among hosts and body sites, with implications for understanding host-microbial dynamics and human-microbial coevolution.

Aggression, rank and power: why hens (and other animals) do not always peck according to their strength.

Thorlief Schjelderup-Ebbe's seminal paper on the 'pecking' order of chickens inspired numerous ethologists to research and debate the phenomenon of dominance. The expansion of dominance to the broader concept of power facilitated disentangling aggression, strength, rank and power. Aggression is only one means of coercing other individuals, and can sometimes highlight a lack of power. The fitness advantages of aggression may only outweigh the costs during periods of uncertainty. Effective instruments of power also include incentives and refusals to act. Moreover, the stability of the power relationship might vary with the instruments used if different means of power vary in the number and types of outcomes achieved, as well as the speed of accomplishing those outcomes. In well-established relationships, actions or physiological responses in the subordinate individual may even be the only indicator of a power differential. A focus on strength, aggression and fighting provides an incomplete understanding of the power landscape that individuals actually experience. Multiple methods for constructing hierarchies exist but greater attention to the implications of the types of data used in these constructions is needed. Many shifts in our understanding of power were foreshadowed in Schjelderup-Ebbe's discussion about deviations from the linear hierarchy in chickens. This article is part of the theme issue 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies'.

Social groups constrain the spatiotemporal dynamics of wild sifaka gut microbiomes.

Primates acquire gut microbiota from conspecifics through direct social contact and shared environmental exposures. Host behaviour is a prominent force in structuring gut microbial communities, yet the extent to which group or individual-level forces shape the long-term dynamics of gut microbiota is poorly understood. We investigated the effects of three aspects of host sociality (social groupings, dyadic interactions, and individual dispersal between groups) on gut microbiome composition and plasticity in 58 wild Verreaux's sifaka (Propithecus verreauxi) from six social groups. Over the course of three dry seasons in a 5-year period, the six social groups maintained distinct gut microbial signatures, with the taxonomic composition of individual communities changing in tandem among coresiding group members. Samples collected from group members during each season were more similar than samples collected from single individuals across different years. In addition, new immigrants and individuals with less stable social ties exhibited elevated rates of microbiome turnover across seasons. Our results suggest that permanent social groupings shape the changing composition of commensal and mutualistic gut microbial communities and thus may be important drivers of health and resilience in wild primate populations.

The long lives of primates and the 'invariant rate of ageing' hypothesis.

Is it possible to slow the rate of ageing, or do biological constraints limit its plasticity? We test the 'invariant rate of ageing' hypothesis, which posits that the rate of ageing is relatively fixed within species, with a collection of 39 human and nonhuman primate datasets across seven genera. We first recapitulate, in nonhuman primates, the highly regular relationship between life expectancy and lifespan equality seen in humans. We next demonstrate that variation in the rate of ageing within genera is orders of magnitude smaller than variation in pre-adult and age-independent mortality. Finally, we demonstrate that changes in the rate of ageing, but not other mortality parameters, produce striking, species-atypical changes in mortality patterns. Our results support the invariant rate of ageing hypothesis, implying biological constraints on how much the human rate of ageing can be slowed.

Female Power: A New Framework for Understanding "Female Dominance" in Lemurs.

Female dominance is often associated with lemurs. However, consensus does not exist among primatologists on how to define, measure, or explain female dominance. This review explores the utility of applying a broader concept of power to understanding lemur intersexual relationships. In this framework, power is defined as arising from an asymmetry in a dyadic relationship and can be divided into 2 types: dominance and leverage. Intersexual asymmetries based upon females having superior fighting ability are considered female dominance. However, economic power also exists, and females with resource-based power exhibit female leverage. Additionally, power has 4 characteristics (base, means, amount, and scope) that describe the precise nature of observed phenomena. This article utilizes the 4 characteristics outlined in the power framework to review the existing "female dominance" literature for lemurs and highlights the value of adopting both an expanded concept of power and a more precise language. By placing the multiple phenomena currently labeled under the single term "female dominance" within the power framework, much of the confusion disappears. Thus, not only is the debate reframed, facilitating endeavors to find evolutionary explanations, but the uniqueness of female power in lemurs can be determined empirically rather than by definition.

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.

Subordination signals improve the quality of social relationships in Verreaux's Sifaka: Implications for the evolution of power structures and social complexity.

OBJECTIVES: Unidirectional dominance-related signals can be used to communicate submission (an immediate behavioral response) or subordination (the status of an established relationship). Subordination signals are defined as emitted during peaceful interactions and are hypothesized to be critical for the evolution of social complexity and robust power structures because they reduce uncertainty in social relationships. The chatter vocalization in Verreaux's sifaka (Propithecus verreauxi) is a unidirectional submissive signal. I tested the hypothesis that chatter vocalizations can signal subordination and thereby reduce agonism in a dyad. MATERIALS AND METHODS: I examined 780 chatters from 18 dyads collected over 881 observation hours on four groups of sifaka in Kirindy Forest, Madagascar. RESULTS: Sifaka emitted 63% of chatters in the peaceful context. Peaceful chatters significantly predicted grooming rate, fighting rate, reconciliation, and proportion of wins in a dyad but did not predict time in proximity. Dyad-type significantly predicted the frequency of peaceful chatters, with intrasexual dyads exhibiting chatters in peaceful contexts more often than intersexual dyads. DISCUSSION: Sifaka communicate both submission and subordination with chatter vocalizations. Subordination signaling increased tolerance and affiliation. It reduced conflicts and the probability dominant individuals usurped resources. Moreover, intrasexual power may be more institutionalized than intersexual power in sifaka. The finding of complex and cognitively demanding social communication in a lemur with low levels of cooperation (1) challenges previous assumptions that the evolution of social complexity is dependent on frequent triadic interactions and high levels of cooperation, and (2) highlights the need for taxonomic diversity in studies of social complexity.

Terrestriality and bacterial transfer: a comparative study of gut microbiomes in sympatric Malagasy mammals.

The gut microbiomes of mammals appear to mirror their hosts' phylogeny, suggesting host-driven microbial community assembly. Yet, much of this evidence stems from comparative studies of distinct wild or captive populations that lack data for disentangling the relative influences of shared phylogeny and environment. Here, we present phylogenetic and multivariate analyses of gut microbiomes from six sympatric (i.e., co-occurring) mammal species inhabiting a 1-km2 area in western Madagascar-three lemur and three non-primate species-that consider genetic, dietary, and ecological predictors of microbiome functionality and composition. Host evolutionary history, indeed, appears to shape gut microbial patterns among both closely and distantly related species. However, we also find that diet-reliance on leaves versus fruit-is the best predictor of microbiome similarity among closely related lemur species, and that host substrate use-ground versus tree-constrains horizontal transmission via incidental contact with feces, with arboreal species harboring far more distinct communities than those of their terrestrial and semi-terrestrial counterparts.

Hierarchical social networks shape gut microbial composition in wild Verreaux's sifaka.

In wild primates, social behaviour influences exposure to environmentally acquired and directly transmitted microorganisms. Prior studies indicate that gut microbiota reflect pairwise social interactions among chimpanzee and baboon hosts. Here, we demonstrate that higher-order social network structure-beyond just pairwise interactions-drives gut bacterial composition in wild lemurs, which live in smaller and more cohesive groups than previously studied anthropoid species. Using 16S rRNA gene sequencing and social network analysis of grooming contacts, we estimate the relative impacts of hierarchical (i.e. multilevel) social structure, individual demographic traits, diet, scent-marking, and habitat overlap on bacteria acquisition in a wild population of Verreaux's sifaka (Propithecus verreauxi) consisting of seven social groups. We show that social group membership is clearly reflected in the microbiomes of individual sifaka, and that social groups with denser grooming networks have more homogeneous gut microbial compositions. Within social groups, adults, more gregarious individuals, and individuals that scent-mark frequently harbour the greatest microbial diversity. Thus, the community structure of wild lemurs governs symbiotic relationships by constraining transmission between hosts and partitioning environmental exposure to microorganisms. This social cultivation of mutualistic gut flora may be an evolutionary benefit of tight-knit group living.

Characterization of the Tree Holes Used by Lepilemur ruficaudatus in the Dry, Deciduous Forest of Kirindy Mitea National Park.

Knowledge of the key resources for a species is critical for developing an effective conservation strategy. Kirindy Mitea National Park is an important refuge for the red-tailed sportive lemur (Lepilemur ruficaudatus), a nocturnal folivorous lemur endemic to the dry deciduous forest of western Madagascar. Because L. ruficaudatus sleeps in tree holes during the day, sleeping trees may be an important resource for this species. Our goal was to characterize the sleeping sites used by L. ruficaudatus at the Ankoatsifaka Research Station in Kirindy Mitea National Park. In July and August 2012, tree characteristics were recorded for 60 L. ruficaudatus sleeping sites. Intact and alive trees, particularly Strychnos madagascariensis, were preferred by L. ruficaudatus. Sleeping holes were generally located in taller trees (median hole height = 4 m) and in trees with large girths (median = 20.5 cm). Greater protection from predators and thermal shifts may be provided by intact trees and concealed tree holes. Because tree characteristics can influence the presence and abundance of forest-living primates, the availability and characteristics of particular trees as potential sleeping shelters in a habitat must be taken into account in conservation strategies for L. ruficaudatus.

Patterns of male dispersal in Verreaux's sifaka (Propithecus verreauxi) at Kirindy Mitea National Park.

Males of many group-living mammals emigrate from their social groups to improve mating opportunities. To help mitigate the social and locational costs of dispersal and to increase reproductive benefits, they may immigrate seasonally, immigrate alongside a partner, and/or replace the alpha male (versus entering a group as a subordinate). Verreaux's sifaka (Propithecus verreauxi) are highly seasonally breeding primates characterized by male-biased dispersal. We hypothesized that (i) males time immigrations to pursue immediate mating opportunities, (ii) entrances with partners more often result in alpha male replacement, and (iii) male competitive ability affects immigration strategy and alpha male tenure length. To assess these proximate aspects of male dispersal, we examined 7 years of demographic, morphological, and behavioral data for five social groups of Verreaux's sifaka in the Kirindy Mitea National Park in western Madagascar. Contrary to expectations and studies of sifaka dispersal in other populations, we detected no strong seasonal pattern in immigrations. Immigrations occurred individually and with partners, and a trend existed for partners to increase the likelihood of replacing an alpha male. Pronounced activity of the sternal scent gland (a proxy for testosterone and prior dominance status), but not body mass, canine size, or potential correlates of leaping ability, significantly influenced immigration strategy. The absence of a seasonal immigration pattern suggests that fluid group boundaries may allow mating success without establishment in a social group before the mating season. Our results also suggest that male immigration strategies are affected by age, prior dominance status, and testosterone levels but not morphological indicators of individual competitive ability. Coalitions may be used to improve competitive ability. Finally, differences in seasonal immigration patterns and length of alpha male vacancies observed at Kirindy Mitea may be due to the relatively low population density. Am. J. Primatol. 79:e22455, 2017. © 2015 Wiley Periodicals, Inc.

Group benefit associated with polymorphic trichromacy in a Malagasy primate (Propithecus verreauxi).

In some primate lineages, polymorphisms in the X-linked M/LWS opsin gene have produced intraspecific variation in color vision. In these species, heterozygous females exhibit trichromacy, while males and homozygous females exhibit dichromacy. The evolutionary persistence of these polymorphisms suggests that balancing selection maintains color vision variation, possibly through a 'trichromat advantage' in detecting yellow/orange/red foods against foliage. We identified genetic evidence of polymorphic trichromacy in a population of Verreaux's sifaka (Propithecus verreauxi) at Kirindy Mitea National Park in Madagascar, and explored effects of color vision on reproductive success and feeding behavior using nine years of morphological, demographic, and feeding data. We found that trichromats and dichromats residing in social groups with trichromats exhibit higher body mass indices than individuals in dichromat-only groups. Additionally, individuals in a trichromat social group devoted significantly more time to fruit feeding and had longer fruit feeding bouts than individuals in dichromat-only groups. We hypothesize that, due to small, cohesive sifaka social groups, a trichromat advantage in detecting productive fruit patches during the energetically stressful dry season also benefits dichromats in a trichromat's group. Our results offer the first support for the 'mutual benefit of association' hypothesis regarding the maintenance of polymorphic trichromacy in primates.

Bidirectional Counterregulation of Human Lung Mast Cell and Airway Smooth Muscle ß2 Adrenoceptors.

Human lung mast cells (HLMCs) play a central role in asthma pathogenesis through their relocation to the airway smooth muscle (ASM) bundles. ß2 adrenoceptor (ß2-AR)-agonists are used to relieve bronchoconstriction in asthma, but may reduce asthma control, particularly when used as monotherapy. We hypothesized that HLMC and human ASM cell (HASMC) responsiveness to ß2-AR agonists would be attenuated when HLMCs are in contact with HASMCs. Cells were cultured in the presence of the short-acting ß2-agonist albuterol, and the long-acting ß2-agonists formoterol and olodaterol. Constitutive and FcεRI-dependent HLMC histamine release, HASMC contraction, and ß2-AR phosphorylation at Tyr(350) were assessed. Constitutive HLMC histamine release was increased in HLMC-HASMC coculture and this was enhanced by ß2-AR agonists. Inhibition of FcεRI-dependent HLMC mediator release by ß2-agonists was greatly reduced in HLMC-HASMC coculture. These effects were reversed by neutralization of stem cell factor (SCF) or cell adhesion molecule 1 (CADM1). ß2-AR agonists did not prevent HASMC contraction when HLMCs were present, but this was reversed by fluticasone. ß2-AR phosphorylation at Tyr(350) occurred within 5 min in both HLMCs and HASMCs when the cells were cocultured, and was inhibited by neutralizing SCF or CADM1. HLMC interactions with HASMCs via CADM1 and Kit inhibit the potentially beneficial effects of ß2-AR agonists on these cells via phosphorylation of the ß2-AR. These results may explain the potentially adverse effects of ß2-ARs agonists when used for asthma therapy. Targeting SCF and CADM1 may enhance ß2-AR efficacy, particularly in corticosteroid-resistant patients.

Biomedical evaluation of Verreaux's sifaka (Propithecus verreauxi) from Kirindy Mitea National Park in Madagascar.

Evaluation of the health, prevalence, and incidence of diseases in wild populations is a critical component of wildlife management. In addition, the establishment of reference physiologic parameters can Verreaux's sifaka (Propithecus verreauxi) from Kirindy Mitea National Park, Madagascar, during two field captures incontribute to the assessment of population risks. Complete medical evaluations were performed on 33 wild June 2010 and June 2011. Each animal received a complete physical examination: weight, body temperature, heart rate, and respiratory rate were recorded, and ectoparasites collected. Blood samples were collected for complete blood cell count, differential white blood cell count, hemoparasite examination, serum biochemical profile, fat-soluble vitamin analysis, trace mineral analysis, and toxoplasmosis and viral serology. Fecal samples were collected for bacterial culture and endoparasite examination. Significant differences exist between age classes for neutrophil and lymphocyte count, alkaline phosphatase, and creatinine values. Parasites detected were Callistoura sp., Bertiella lemuriformis, and Dipetalonema petteri. This publication reports the first complete biomedical evaluation of the P. verreauxi and provides a basis for hematologic and biochemical comparisons of P. verreauxi in the wild.

Noninvasive molecular sexing: an evaluation and validation of the SRY- and amelogenin-based method in three new lemur species.

Many lemur species are arboreal, elusive, and/or nocturnal and are consequently difficult to approach, observe and catch. In addition, most of them are endangered. For these reasons, non-invasive sampling is especially useful in primates including lemurs. A key issue in conservation and ecological studies is to identify the sex of the sampled individuals to investigate sex-biased dispersal, parentage, social organization and population sex ratio. Several molecular tests of sex are available in apes and monkeys, but only a handful of them work in the lemuriform clade. Among these tests, the coamplification of the SRY gene with the amelogenin X gene using strepsirhine-specific X primers seems particularly promising, but the reliability and validity of this sexing test have not been properly assessed yet. In this study, we (i) show that this molecular sexing test works on three additional lemur species (Microcebus tavaratra, Propithecus coronatus and P. verreauxi) from two previously untested genera and one previously untested family, suggesting that these markers are likely to be universal among lemurs and other strepsirrhines; (ii) provide the first evidence that this PCR-based sexing test works on degraded DNA obtained from noninvasive samples; (iii) validate the approach using a large number of known-sex individuals and a multiple-tubes approach, and show that mismatches between the field sex and the final molecular consensus sex occur in less than 10% of all the samples and that most of these mismatches were likely linked to incorrect sex determinations in the field rather than genotyping errors.

Effects of the distribution of female primates on the number of males.

The spatiotemporal distribution of females is thought to drive variation in mating systems, and hence plays a central role in understanding animal behavior, ecology and evolution. Previous research has focused on investigating the links between female spatiotemporal distribution and the number of males in haplorhine primates. However, important questions remain concerning the importance of spatial cohesion, the generality of the pattern across haplorhine and strepsirrhine primates, and the consistency of previous findings given phylogenetic uncertainty. To address these issues, we examined how the spatiotemporal distribution of females influences the number of males in primate groups using an expanded comparative dataset and recent advances in bayesian phylogenetic and statistical methods. Specifically, we investigated the effect of female distributional factors (female number, spatial cohesion, estrous synchrony, breeding season duration and breeding seasonality) on the number of males in primate groups. Using bayesian approaches to control for uncertainty in phylogeny and the model of trait evolution, we found that the number of females exerted a strong influence on the number of males in primate groups. In a multiple regression model that controlled for female number, we found support for temporal effects, particularly involving female estrous synchrony: the number of males increases when females are more synchronously receptive. Similarly, the number of males increases in species with shorter birth seasons, suggesting that greater breeding seasonality makes defense of females more difficult for male primates. When comparing primate suborders, we found only weak evidence for differences in traits between haplorhines and strepsirrhines, and including suborder in the statistical models did not affect our conclusions or give compelling evidence for different effects in haplorhines and strepsirrhines. Collectively, these results demonstrate that male monopolization is driven primarily by the number of females in groups, and secondarily by synchrony of female reproduction within groups.