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.

The Influence of Seasonal Availability of Young Leaves on Dietary Niche Separation in Two Ecologically Similar Folivorous Lemurs.

Traditional socio-ecological models consider that folivorous primates experience limited feeding competition due to the low quality, high abundance, and even distribution of leaves. Evidence from several folivorous species that experience similar constraints to frugivores does not support this hypothesis. The sympatric lemur genera Avahi (Indriidae) and Lepilemur (Lepilemuridae) are good models to understand how food availability constrains folivores since they are both nocturnal, folivorous, and have a comparable body mass. Here we investigate how two nocturnal folivorous primates, Avahi meridionalis and Lepilemur fleuretae, living in the lowland rain forest of Tsitongambarika, South-East Madagascar, partition their dietary niche and are influenced by seasonality of young leaves. To account for food availability, we collected annual phenological data on 769 trees from 200 species. We also collected behavioural data on 5 individuals per lemur species from August 2015 to July 2016 via continuous focal sampling. We found the phenological profile to be seasonal with peaks of leaf flushing, flowering, and fruiting occurring in the austral summer. The two species showed limited dietary overlap (37% rich period, 6% lean period), and A. meridionalis showed higher feeding time and longer daily distances travelled during the rich period. Lepilemur fleuretae showed a dietary shift during the lean period, relying more on mature leaves (73.3% during the lean period, 13.5% during the rich period) but maintaining similar activity levels between seasons. The time spent feeding on food items by A. meridionalis was positively correlated with the nitrogen content and negatively correlated with polyphenols during the rich period. We highlighted a clear effect of the seasonality of young leaves on the diet, nutritional content, activity patterns, and daily distances travelled by two folivorous species, which can be linked to nutrient balancing and time-minimising versus energy-maximising strategies.

Mitogenomes of historical type specimens unravel the taxonomy of sportive lemurs (Lepilemur spp.) in Northwest Madagascar.

Accurate information on name-bearing types, including corresponding type localities, is essential for proper taxonomy. However, such geographic information is often missing or unreliable. The localities of type specimens collected 100-200 years ago can be difficult to trace due to changes in local names or simple inaccuracies. Such a case can be found for the gray-backed sportive lemur (Lepilemur dorsalis), with its type locality imprecisely fixed as Northwest Madagascar. In recent years, eight species have been newly described for the Inter-River-Systems (IRSs) of this region, however the designation of L. dorsalis remains controversial due to a lack of a precise type locality. Here, we sequenced the complete mitochondrial genomes (mitogenomes) of type specimens of L. dorsalis and L. grandidieri, which is currently recognized as a synonym of L. dorsalis and compared their sequences with those of samples of known provenance from different IRSs. Results showed that the two type specimens of L. dorsalis and L. grandidieri had identical mitogenome sequences and clustered closely with samples collected in IRS V, indicating that the type locality could be fixed to IRS V. Consequently, L. dorsalis occurs in IRS V, and L. grandidieri and L. mittermeieri are junior synonyms of L. dorsalis. This finding demonstrates the value of type specimens for clarifying phylogeographic and taxonomic questions and clarifies the taxonomy of sportive lemurs in Northwest Madagascar.

The role of developmental rate, body size, and positional behavior in the evolution of covariation and evolvability in the cranium of strepsirrhines and catarrhines.

Recent studies on hominin craniofacial evolution have focused on phenotypic integration or covariation among traits. Covariation is thought to significantly affect evolutionary trajectories, shaping the ways in which hominins and other primates could have evolved. However, the ways in which covariation itself evolves are not well understood. This study aims to investigate the role of phylogeny, development, body size, and positional behavior in shaping the strength of covariation in strepsirrhine and catarrhine primate crania (n = 1009, representing 11 genera). These factors may have been catalysts for change in the magnitude of covariation, and they have changed significantly during primate evolution and particularly hominin evolution. Modern humans in particular have slow developmental trajectories, large bodies, and a unique form of locomotion in the form of orthograde bipedalism. Variance of eigenvalues, mean integration, mean evolvability, and mean conditional evolvability was estimated and their relationship to the various factors described earlier was assessed using phylogenetic and nonphylogenetic analyses. Results indicate that some phylogenetic signal is present, but it is not equivalent across integration statistics or cranial regions. In particular, these results suggest that closely related species are more similar than more distantly related species in evolvability of the cranial base and integration of the face. Two divergent patterns were also identified, in which covariation and evolvability of the cranial base are linked to developmental rate, but those of the face are linked to body size. Neither locomotion nor posture appears related to covariation or evolvability of the primate cranium. These results suggest that overall low covariation observed in the hominin cranium may be a result of separate trends in different cranial regions.

The acoustic near-field measurement of aye-ayes' biological auditory system utilizing a biomimetic robotic tap-scanning.

The aye-aye (Daubentonia madagascariensis) is best known for its unique acoustic-based foraging behavior called 'tap-scanning' or 'percussive foraging'. The tap-scanning is a unique behavior allowing aye-aye to locate small cavities beneath tree bark and extract wood-boring larvae from it. The tap-scanning requires the animal auditory system to have exceptional acoustic near-field sensitivity. This paper has experimentally investigated the effects of external pinna in the acoustic sensing and detection capabilities of aye-ayes. To experimentally evaluate the effects of external ear (pinna) of the aye-aye, the tap-scanning process was simulated using a robotic arm. A pinna was 3D printed using a CT scan obtained from a carcass. The pinna's effect on the acoustic near-field has been evaluated in time and frequency domains for simulated tap-scanning with the pinna in upright and cupped positions. This idea originates from behavioral observations of the aye-aye using its ears in this way. The results suggest that the aye-aye can substantially enhance its acoustic near-field sensitivity through a cupped conformation during tap-scanning. Three phenomena contribute to this substantial enhancement of the acoustic near-field: (i) a considerable increase in the signal-to-noise ratio, (ii) the creation of a focal area and potentially a focal point to increase the spatial resolution, and (iii) an increase in the receiver peak frequency by changing near-field beam pattern for higher frequencies that can result in greater sensitivity due to a smaller wavelength.

Design, delivery and perception of condition-dependent chemical signals in strepsirrhine primates: implications for human olfactory communication.

The study of human chemical communication benefits from comparative perspectives that relate humans, conceptually and empirically, to other primates. All major primate groups rely on intraspecific chemosignals, but strepsirrhines present the greatest diversity and specialization, providing a rich framework for examining design, delivery and perception. Strepsirrhines actively scent mark, possess a functional vomeronasal organ, investigate scents via olfactory and gustatory means, and are exquisitely sensitive to chemically encoded messages. Variation in delivery, scent mixing and multimodality alters signal detection, longevity and intended audience. Based on an integrative, 19-species review, the main scent source used (excretory versus glandular) differentiates nocturnal from diurnal or cathemeral species, reflecting differing socioecological demands and evolutionary trajectories. Condition-dependent signals reflect immutable (species, sex, identity, genetic diversity, immunity and kinship) and transient (health, social status, reproductive state and breeding history) traits, consistent with socio-reproductive functions. Sex reversals in glandular elaboration, marking rates or chemical richness in female-dominant species implicate sexual selection of olfactory ornaments in both sexes. Whereas some compounds may be endogenously produced and modified (e.g. via hormones), microbial analyses of different odorants support the fermentation hypothesis of bacterial contribution. The intimate contexts of information transfer and varied functions provide important parallels applicable to olfactory communication in humans. This article is part of the Theo Murphy meeting issue 'Olfactory communication in humans'.

First navicular remains of a European adapiform (Anchomomys frontanyensis) from the Middle Eocene of the Eastern Pyrenees (Catalonia, Spain): implications for early primate locomotor behavior and navicular evolution.

We describe the first known navicular bones for an Eocene euprimate from Europe and assess their implications for early patterns of locomotor evolution in primates. Recovered from the fossil site of Sant Jaume de Frontanyà-3C (Barcelona, Spain), the naviculars are attributed to Anchomomys frontanyensis. The small size of A. frontanyensis allows us to consider behavioral implications of comparisons with omomyiforms, regardless of allometric sources of navicular variation. Researchers usually consider omomyiforms to be more prone to leaping than contemporaneous adapiforms partly because of the more pronounced elongation of omomyiform tarsal elements. However, A. frontanyensis differs from other adapiforms and is similar to some omomyiforms in its more elongated navicular proportions. Although this might raise questions about attribution of these naviculars to A. frontanyensis, the elements exhibit clear strepsirrhine affinities leaving little doubt about the attribution: the bones' mesocuneiform facets contact their cuboid facets. We further propose that this strepsirrhine-specific feature in A. frontanyensis and other adapiforms reflects use of more inverted foot postures and potentially smaller substrates than sympatric omomyiforms that lack it. Thus substrate differences may have influenced niche partitioning in Eocene euprimate communities along with differences in locomotor agility. As previous studies on the astragalus and the calcaneus have suggested, this study on the navicular is consistent with the hypothesis that the locomotor mode of A. frontanyensis was similar to that of extant cheirogaleids, especially species of Microcebus and Mirza.

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.

Re-assessing the applicability of the Jarman/Bell model and Kay's threshold to the insectivorous aye-aye (Daubentonia madagascariensis).

OBJECTIVES: Jarman/Bell principle and Kay's threshold suggest that large animals should not be able to sustain themselves on insects. However, animals with specialized morphological and/or behavioral adaptations violate these assumptions. Male aye-ayes were recently identified as having an insectivorous diet despite weighing 2.5 kg. We further explored this diet/body size disparity by studying behaviors of an adult female and juvenile male aye-aye. MATERIALS AND METHODS: We collected behavioral data on an adult female and juvenile male aye-aye in Torotorofotsy, Madagascar from January 2016 to December 2017. We used instantaneous sampling to determine the frequency of feeding events and continuous sampling during feeding behaviors to assess duration of feeding bouts. RESULTS: Invertebrates comprised over 88% of the diet for both animals. A Wilcoxon signed-rank test found no significant difference in the female's feeding frequencies between the hot/rainy seasons and the cold/dry seasons. DISCUSSION: Our results support earlier findings that invertebrates are the aye-aye's main resource and corroborate that aye-ayes violate assumptions of the Jarman/Bell model and Kay's threshold. We suggest that the Jarman/Bell principle and Kay's threshold be used as supporting evidence and not to repudiate findings that do not conform to these guidelines.

DiceCT Analysis of the Extreme Gouging Adaptations Within the Masticatory Apparatus of the Aye-Aye (Daubentonia madagascariensis).

Relative to all other primates, the aye-aye (Daubentonia madagascariensis) exists at the extremes of both morphology and behavior. Its specialized anatomy-which includes hypselodont incisors and highly derived manual digits-reflects a dietary niche, unique among primates, which combines tap-foraging with gouging to locate and extract wood-boring larvae. Here, we explore the impact of this extreme dietary ecology upon the masticatory musculature of this taxon with reference to a second, similarly sized but highly generalist lemuriform-the mongoose lemur (Eulemur mongoz). Using non-destructive, high-resolution diffusible iodine-based contrast-enhanced computed tomography techniques, we reconstruct the three-dimensional volumes of eight masticatory muscles, and, for the first time in strepsirrhines, isolate and visualize their constituent muscle fascicles in situ and in three dimensions. Using these data, we report muscle volumes, forces, and fascicle lengths from each muscle portion, as well as their orientation relative to two standardized anatomical planes. Our findings demonstrate the overbuilt nature of the aye-aye's masticatory apparatus, in which each muscle possesses an absolutely and relatively larger muscle volume and PCSA than its counterpart in the mongoose lemur. Likewise, for several adductor muscles, aye-ayes also possess relatively greater fascicle lengths. Finally, we note several unusual features within the lateral pterygoid of the aye-aye-the muscle most responsible for jaw protrusion-that relate to force maximization and reorientation. As this jaw motion is critical to gouging, we interpret these differences to reflect highly specific specializations that facilitate the aye-aye's extreme subsistence strategy. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:282-294, 2020. © 2019 American Association for Anatomy.

Nocturnal Primate Communication: Ecology, Evolution and Conservation.

Auditory, visual and olfactory cues play varying roles in non-human primate communication, and these systems have been intensively studied over the last several decades. The use of vocalisations as a primary mode of communication has been the focus of much research, especially in attempts to understand the origins of human language, with a major focus on anthropoid primates and diurnal lemurs. Over the last decade, technological advances have allowed researchers to begin to conduct in-depth investigations into the communication systems exhibited by the nocturnal and cathemeral prosimian primates, including tarsiers, lemurs and lorises. Understanding how nocturnal prosimians use visual, olfactory and auditory cues is vital for reconstructing the origins of primate communication systems. In this special issue, we highlight some of the more exciting advances in the communication strategies of the prosimians. Contributions come from work conducted in Kenya, Tanzania, Senegal, Rwanda, Madagascar and the Indonesian islands of Java and Sulawesi. Topics will include: the description of novel ultrasonic vocalisations, including frequency and function of these newly discovered calls; the possible use of vocalisations to navigate and assemble at sleep sites; the importance of species-specific contact vocalisations for the identification of new species; the use of urinary and glandular signals to communicate and the methods developed to understand this complex communication in the field; the use of vocalisations for niche separation among nocturnal primates from mainland Africa and Madagascar; and whether or not we can use new technologies to discern whether prosimians use vocalisations for individual identification of group members. We discuss the importance of new field methods including novel equipment and techniques, the use of vocalisation to influence conservation practices and the importance of comparing across prosimian taxa to reconstruct the communication systems of our early primate ancestors.

P-Mail: The Information Highway of Nocturnal, but Not Diurnal or Cathemeral, Strepsirrhines.

Scent marking is a well-established, but highly variable, mode of communication among strepsirrhine primates. We begin by reviewing this literature, focusing on nocturnal species. Our understanding about the information content of scent signals and the factors driving species diversity remains incomplete, owing to difficulties in acquiring comparative chemical data. We therefore re-examine such a data set, representing the richness and relative abundance of volatile organic compounds (VOCs) in the urine of 12 species (from Galagidae, Lorisidae, Daubentoniidae, Cheirogaleidae, Indriidae, and Lemuridae), to explore differences between nocturnal, diurnal and cathemeral species. As predicted by the variable importance of urine marking across species, the urine of nocturnal strepsirrhines contained the most VOCs and putative semiochemicals, differed significantly in composition from that of diurnal and cathemeral species and showed the strongest species scent "signatures." Relevant to retracing the evolutionary trajectory of cathemeral strepsirrhines, nocturnal and diurnal species were most differentiated in their VOCs, with cathemeral species being intermediary, but more closely aligned with diurnal species. These data support cathemerality as an ancient expansion of diurnal animals into a nocturnal niche. Consideration of the traits and variables associated with olfactory communication offers a profitable new way for examining species diversity and patterns of evolutionary change.

Sahamalaza Sportive Lemur, Lepilemur sahamalaza, Vocal Communication: Call Use, Context and Gradation.

Stereotypical vocalisations can facilitate long-distance communication in dense and, thus, sound-degrading forest habitats. Despite this, primate vocal repertoires often also include gradations between different call types that are used in social interactions. Because many nocturnal primates show a solitary social structure, it has been difficult to assess the role vocalisations play in mediating their social encounters. Here we aim to expand on the call types of Sahamalaza sportive lemurs, Lepilemur sahamalaza, as well as investigate their variability and use in social contexts. Through long-term behavioural observations with concurrent recording of vocalisations of known and unknown individuals between 2015 and 2016, we described three previously unrecorded calls and expanded the vocal repertoire to 12 different call types that were used in both targeted social and broadcast solitary contexts. Linear discriminant function and cluster analysis supported initial observations that vocalisations within call types were variable and that at least two call types can be classified as "graded," contrary to expectations. Despite variations in call-context connections, no clear patterns of call use could be established. However, measurable differences in vocal behaviour between the seasons and the sexes indicate that calling is used in a reproductive context, similar to other nocturnal and diurnal primates.

Eulerian videography technology improves classification of sleep architecture in primates.

Sleep is a critically important dimension of primate behavior, ecology, and evolution, yet primate sleep is under-studied because current methods of analyzing sleep are expensive, invasive, and time-consuming. In contrast to electroencephalography (EEG) and actigraphy, videography is a cost-effective and non-invasive method to study sleep architecture in animals. With video data, however, it is challenging to score subtle changes that occur in different sleep states, and technology has lagged behind innovations in EEG and actigraphy. Here, we applied Eulerian videography to magnify pixels relevant to scoring sleep from video, and then compared these results to analyses based on actigraphy and standard infrared videography. We studied four species of lemurs (Eulemur coronatus, Lemur catta, Propithecus coquereli, Varecia rubra) for 12-h periods per night, resulting in 6480 1-min epochs for analysis. Cramer's V correlation between actigraphy-classified sleep and infrared videography-classified sleep revealed consistent results in eight of the nine 12-h videos scored. A sample of the infrared videography was then processed by Eulerian videography for movement magnification and re-coded. A second Cramer's V correlation analysis, between two independent scorers coding the same Eulerian-processed video, found that interobserver agreement among Eulerian videography increased sleep vs. awake, NREM, and REM classifications by 7.1%, 46.7%, and 34.3%, respectively. Furthermore, Eulerian videography was more strongly correlated with actigraphy data when compared to results from standard infrared videography. The increase in agreement between the two scorers indicates that Eulerian videography has the potential to improve the identification of sleep states in lemurs and other primates, and thus to expand our understanding of sleep architecture without the need for EEG.

A comparative study of gut microbiomes in captive nocturnal strepsirrhines.

Feeding strategy and diet are increasingly recognized for their roles in governing primate gut microbiome (GMB) composition. Whereas feeding strategy reflects evolutionary adaptations to a host's environment, diet is a more proximate measure of food intake. Host phylogeny, which is intertwined with feeding strategy, is an additional, and often confounding factor that shapes GMBs across host lineages. Nocturnal strepsirrhines are an intriguing and underutilized group in which to examine the links between these three factors and GMB composition. Here, we compare GMB composition in four species of captive, nocturnal strepsirrhines with varying feeding strategies and phylogenetic relationships, but nearly identical diets. We use 16S rRNA sequences to determine gut bacterial composition. Despite similar husbandry conditions, including diet, we find that GMB composition varies significantly across host species and is linked to host feeding strategy and phylogeny. The GMBs of the omnivorous and the frugivorous species were significantly more diverse than were those of the insectivorous and exudativorous species. Across all hosts, GMBs were enriched for bacterial taxa associated with the macronutrient resources linked to the host's respective feeding strategy. Ultimately, the reported variation in microbiome composition suggests that the impacts of captivity and concurrent diet do not overshadow patterns of feeding strategy and phylogeny. As our understanding of primate GMBs progresses, populations of captive primates can provide insight into the evolution of host-microbe relationships, as well as inform future captive management protocols that enhance primate health and conservation.

Ambient occlusion and PCV (portion de ciel visible): A new dental topographic metric and proxy of morphological wear resistance.

Recently, ambient occlusion, quantified through portion de ciel visible (PCV) was introduced as a method for quantifying dental morphological wear resistance and reconstructing diet in mammals. Despite being used to reconstruct diet and investigate the relationship between dental form and function, no rigorous analysis has investigated the correlation between PCV and diet. Using a sample of platyrrhine and prosimians M2s, we show average PCV was significantly different between most dietary groups. In prosimian, insectivores had the lowest PCV, followed by folivores, omnivores, frugivores, and finally hard-object feeders. In platyrrhines, omnivores had the lowest average PCV, followed by folivores, frugivores, and finally hard-object feeders. PCV was correlated to two topographic variables (Dirichlet normal energy, DNE, and relief index, RFI) but uncorrelated to three others (orientation patch count rotated, OPCR, tooth surface area, and tooth size). The OPCR values here differed greatly from previously published values using the same sample, showing how differences in data acquisition (i.e., using 2.5D vs. 3D surfaces) can lead to drastic differences in results. Compared to other popular topographic variables, PCV performed as well or better at predicting diet in these groups, and when combined with a metric for size, the percent of successful dietary classifications reached 90%. Further, using an ontogenetic series of hominin (Paranthropus robustus) M2s, we show that PCV correlates well with probability of wear, with PCV values being higher on the portions of the occlusal surface that experience more wear (e.g., cusps and crest tips, wear facets) than the portions of the tooth that experience less. This relationship is strongest once wear facets have begun to form on the occlusal surface. These results highlight the usefulness of PCV in quantifying morphological wear resistance and predicting diet in mammals.

Middle ear pneumatization in nonhuman primates: A comparative analysis.

OBJECTIVES: We test the effects of body mass and phylogeny on middle ear cavity pneumatization, and the role of pneumatization in hearing function, spanning the anatomical, ecological, and behavioral diversity of nonhuman primates. MATERIALS AND METHODS: All cavities were segmented in middle ear scans of 96 specimens, from 12 strepsirrhine and 15 haplorhine extant species. We measured the tympanic cavity (TC) separately, and all other middle ear spaces together (MES), calculating the degree of pneumatization with the surface area-to-volume ratio. We tested body mass effect with linear regression; we evaluated the phylogenetic signal and selection patterns, using a Kappa statistic test, and Ornstein-Uhlenbeck models (OU). We investigated the link between pneumatization and hearing sensitivity using phylogenetic regression. RESULTS: Testing body mass reveals an allometric pattern for both TC and MES dimensions. Degree of pneumatization in MES is dependent on body mass in haplorhines: larger animals have more pneumatized MES. Differences at various taxonomic ranks were observed for MES, while no phylogenetic influence was observed for TC. Infraorder selection patterns are different. Auditory performance is significantly related to degree of pneumatization, indicating that a pneumatized middle ear is associated with better perception of low frequencies. DISCUSSION: Pneumatization in MES is under differential selective pressure, indicating several optima for this trait. Pneumatization in MES probably modifies hearing sensitivity through pressure regulation mechanisms, auditory bulla size reduction, and frequency modulation. This could explain strepsirrhine adaptation to high-frequency perception, while haplorhine auditory perception is adapted to a broader sound range, including high and low frequencies.

Spatiotemporal distribution of individuals as an indicator for the social system of Lepilemur sahamalaza.

Primate social systems are highly diverse, complicating the classification of particularly elusive species that are difficult to observe. The spatial distribution of individuals over time is a critica lindicator for the social organization and long-term studies are important to establish patterns of social interactions. In recent years, species of the cryptic, nocturnal sportive lemurs of the genus Lepilemur were found to live in pairs in which a single male and a single female share and defend a mutual home range. The present study aimed to forward research into this underrepresented genus by determining the social organization and structure of the Sahamalaza sportive lemur, L. sahamalaza. We collected 773.15 hr of behavioral and GPS data during a period of 10 months (between 2015 and 2016) on 14 individuals: eight females and six males. There was no evidence of pair-specific home range use as intra- and intersexual home range overlap was high. No pattern of social interactions between focal individuals could be distinguished despite high range overlap. Individuals met and interacted infrequently, resulting in an interaction rate of 0.32 interactions/hr. Sleeping associations between adult individuals were never observed. While both sexes had access to multiple potential mating partners, range sizes or ranging distances did not increase in mating periods. Overall, the social system of Sahamalaza sportive lemurs exhibits aspects of a solitary social organization and structure with potential for the polygamous mating system. These findings underline the importance of detailed social ecology studies that can provide the basis for understanding potential environmental influences on social system variability of closely related species.

Evolutionary dynamics of sexual size dimorphism in non-volant mammals following their independent colonization of Madagascar.

As predicted by sexual selection theory, males are larger than females in most polygynous mammals, but recent studies found that ecology and life history traits also affect sexual size dimorphism (SSD) through evolutionary changes in either male size, female size, or both. The primates of Madagascar (Lemuriformes) represent the largest group of mammals without male-biased SSD. The eco-evo-devo hypothesis posited that adaptations to unusual climatic unpredictability on Madagascar have ultimately reduced SSD in lemurs after dispersing to Madagascar, but data have not been available for comparative tests of the corresponding predictions that SSD is also absent in other terrestrial Malagasy mammals and that patterns of SSD changed following the colonization of Madagascar. We used phylogenetic methods and new body mass data to test these predictions among the four endemic radiations of Malagasy primates, carnivorans, tenrecs, and rodents. In support of our prediction, we found that male-biased SSD is generally absent among all Malagasy mammals. Phylogenetic comparative analyses further indicated that after their independent colonization of Madagascar, SSD decreased in primates and tenrecs, but not in the other lineages or when analyzed across all species. We discuss several mechanisms that may have generated these patterns and conclude that neither the eco-evo-devo hypothesis, founder effects, the island rule nor sexual selection theory alone can provide a compelling explanation for the observed patterns of SSD in Malagasy mammals.

Systematic and locomotor diversification of the Adapis group (Primates, Adapiformes) in the late Eocene of the Quercy (Southwest France), revealed by humeral remains.

Twenty humeral specimens from the old and new Quercy collections attributed to the fossil primates Adapis and Palaeolemur are described and analysed together. We provide a qualitative and quantitative analysis of the different humeri, revealing that high variability is present within the "Adapis group" sample. Six different morphotypes are identified, confirming that what has often been called "Adapis parisiensis" is a mix of different species that present different locomotor adaptations. Such a relatively high level of locomotor diversity is unique in the Paleogene primate fossil record. The humeral proportions of Adapis overlap with different groups of extant strepsirrhines and platyrrhines depending on the specimen, so the popular view of Adapis as a loris-like slow climbing primate does not apply to the whole sample presented here. Moreover, different humeral features traditionally associated with "Adapis parisiensis", such as the absence of a zona conoidea and a reduced brachioradialis flange, are variable depending on the sample studied. In addition, results of our analyses show that adapine and omomyid humeral morphology overlap extensively, leading us to question the accuracy of taxonomic attributions based on morphology of isolated humeri at localities where omomyids and adapines of similar size coexist. Finally, assuming our different morphotypes represent different species within two genera, we propose a phylogenetic hypothesis relating these morphotypes, which inhabited a small geographic area.