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The purpose of this research paper is focusing on the hyoid bone in Alouatta pigra and Alouatta macconnelli and determine its role in affecting the social organization and sexual selection of the individuals in these species. This includes the discussion on the dimensions of the hyoid between the males and females as well as an analysis of observed social behaviour patterns and the sexual selection process. It was found that female Alouatta pigra had a larger hyoid than males of the species and that Alouatta macconnelli males had a hyoid nearly twice the size of females. The hyoids were measured from the Mammal Biodiversity collection at the Royal Ontario Museum. The bones dimensions were taken using plastic calipers and their circumference measured using a soft tape measure. Based on the sexual selection and social organisation of Alouatta pigra which is multi-male/multi-female with the occasional unimale with multiple females, the null hypothesis must be accepted as the larger hyoid is not found in males but in females and the size of the hyoid does not affect the social organization in this species. In Alouatta macconnelli, there is little information on this species but the null hypothesis is rejected as males have much larger hyoids than females and the social organization is that of a uni-male with multiple females.

Studies on the impact of habitat loss on species occurrence consistently find that the amount of habitat (measured as patch area) is a major determinant of species occurrence at a patch-level. However, patch-level research may fail to detect important patterns and processes only observable at a landscape-level. A landscape-level approach that incorporates species-specific scale responses is needed to better understand what drives species occurrence. Our aim was to determine the landscape-level scale of effect of habitat amount on the occurrence of three species of nocturnal lemurs (Cheirogaleus medius, Microcebus murinus, and M. ravelobensis). We surveyed line transects to determine the occurrence of three lemur species within a fragmented landscape of deciduous dry forest and anthropogenic grassland in northwestern Madagascar. To determine the scale of effect of habitat loss on lemur occurrence, we compared logistic regression models of occurrence against habitat amount among eight different landscape scales using Akaike's Information Criterion values. We found differing scale responses among the lemurs in our study. Occurrence of C. medius responded to habitat amount at scales between 0.5-4 ha, M. murinus at scales between 1 and 4 ha and M. ravelobensis at scales between 0.125 and 4 ha. We suggest that the scale of effect for C. medius is mediated by their ability to hibernate. A relatively lower scale-response for Microcebus spp. likely reflect their omnivorous diet, small habitat requirements, and limited dispersal ability. Differences in scale responses between M. murinus and M. ravelobensis are likely a result of differing dispersal ability and responses to edge effects between these species. Our study is among the first on lemurs to show the value of a landscape-level approach when assessing the effects of habitat loss on species occurrence.

Madagascar is home to many threatened and endemic primate species, yet this island has seen dramatic declines in lemur habitat due to forest loss. This forest loss has resulted in an increasingly fragmented forest landscape, with fragments isolated from each other by grasslands (i.e., matrix). The grassland matrix is not entirely homogeneous containing matrix elements such as isolated trees and shrubs and linear features such as drainage lines. Because most lemurs are predominantly arboreal, they may preferentially use matrix elements to facilitate dispersal between fragments for access to mates or reduce feeding competition, allowing gene flow between fragments of habitat. Therefore, it is important to understand to what degree they use the matrix. We investigated matrix use in two mouse lemurs, the grey mouse lemur (Microcebus murinus) and the golden-brown mouse lemur (Microcebus ravelobensis) in a fragmented landscape in northwest Madagascar. We tested the predictions that: (1) lemurs use matrix less often than forest fragments, (2) if they use the matrix, then they will preferentially use matrix elements compared to grassland, and (3) M. murinus will disperse into the matrix further than M. ravelobensis. In 2011, we visually surveyed line transects in four areas containing matrix elements and four adjacent forest fragments during nocturnal walks. In 2017, we set up traplines in four areas of the matrix containing matrix elements, three areas that were grassland, and six traplines in adjacent fragments. We compared the relative abundance of mouse lemurs in matrix transects to fragmented forest transects, and the relative abundance of captured lemurs in matrix elements, grassland, and fragment traplines. We found that encounter rates of mouse lemurs did not significantly differ between the matrix and fragmented forest transects or traplines. Our sample size was too low to determine if the mean distance from the forest was greater for either Microcebus spp. Our study highlights that mouse lemurs do use matrix elements and there may be interspecific differences in use. Further research is needed to confirm species-specific matrix use, why mouse lemurs use matrix, and how much matrix elements facilitate movement for each species in fragmented landscapes.

Habitat loss and fragmentation are major ecological forces threatening animal communities across the globe. These issues are especially true in Madagascar, where forest loss is ongoing. We examined the effects of forest fragmentation on the distribution and abundance of sympatric, endemic gray, and golden-brown mouse lemurs (Microcebus murinus and Microcebus ravelobensis), the endemic western tuft-tailed rat (Eliurus myoxinus), and the invasive black rat (Rattus rattus) in two regions in northwestern Madagascar. We used systematic capture procedures in 40 forest fragments and four continuous forest sites which differed in size, shape, and degree of isolation. With a trapping effort of 11,567 trap nights during two dry seasons (2017-2018), we captured 929 individuals (432 M. ravelobensis, 196 M. murinus, 116 E. myoxinus, and 185 R. rattus). We examined the influence of study region, forest type (fragment vs. continuous), forest size, forest shape, the proportion of 50-m forest edge and distance to continuous forest on the abundance and interaction of the four species. Responses to fragmentation differed strongly between species, but no interaction could be detected between the abundance of the different species. Thus competition within and between native and invasive species may not be regulating abundances in these regions. On the contrary, the abundance of M. ravelobensis and E. myoxinus differed significantly between study regions and was negatively affected by fragmentation. In contrast, there was no evidence of an impact of fragmentation on the abundance of M. murinus. Finally, the invasive R. rattus responded positively to the increasing distance to the continuous forest. In conclusion, the response of small Malagasy mammals to forest fragmentation varies largely between species, and fragmentation effects need to be examined at a species-specific level to fully understand their ecological dynamics and complexity.

Although roads are often assumed to be barriers to the dispersal of arboreal species, there has been little empirical testing of this assumption. If arboreal animals are unable to cross roads, population subdivision may occur, or resources may become inaccessible. We tested the hypothesis that Route Nationale 4 (RN4), a paved highway, was a barrier to movement and dispersal of the Endangered golden-brown mouse lemurMicrocebus ravelobensisin Ankarafantsika National Park, in north-west Madagascar. During June–August 2015 we conducted a capture–mark–recapture study at three sites: two adjacent to RN4 and one within intact forest without a potential barrier. During 2,294 trap nights we captured 120 golden-brown mouse lemurs 1,032 times. In roadside habitats we captured significantly more males than females, whereas the opposite was the case in interior forest habitat. We detected eighteen crossings of highway transects by nine individuals; however, all potential dispersal events involved males. In roadside habitat, movement was significantly inhibited in both males and females. We present some of the first data on the effects of roads on movement patterns in arboreal Malagasy mammals, showing species- and sex-biased effects of roads as dispersal barriers. Our findings indicate that roads may not be complete barriers to dispersal in lemurs. We recommend that conservation managers and scientists examine explicitly the effects of roads and natural arboreal bridges in Madagascar in future studies.

The study of habitat fragmentation is complex because multiple, potentially synergistic, ecological processes may be acting simultaneously. Further, edge effects themselves may be complex in that additivity from multiple edges can give rise to heterogeneous nearest–edge gradients. We used heat diffusion as a proxy for additive edge effects in two study landscapes in order to test whether two key observations recently attributed to synergy between edge and area effects could be more simply explained by additivity; namely, steeper edge gradients in larger fragments and variation in slopes of species–area relationships as a function of distances to fragment edges. We sampled forest structure in northwestern Madagascar at various distances from the edge in fragments and continuous forest and used an inverse modelling approach to parameterize the model. In addition, we applied the model to data from a published study of beetle communities in fragmented forests in New Zealand. With increasing proximity to edges, woody stem densities decreased and, as predicted, smaller fragments had lower stem densities and less steep edge gradients than larger ones. The model successfully predicted shifts in species–area relationships as a function of nearest–edge distances for beetle species, although observed richness for forest specialists in the smallest fragments was lower than predicted. Two key observations attributed to synergy between edge and area effects were explained by edge additivity. The model is particularly useful in that it can help to disentangle the complex sets of processes acting in fragmented landscapes.

We used species-area relationships (SARs) to investigate the effects of habitat loss on lemur biogeography. We measured species richness via visual surveys on line transects within 42 fragments of dry deciduous forest at the Ambanjabe field site in Ankarafantsika National Park, Madagascar. We measured human disturbance and habitat characteristics within 38 of the 42 fragments. We measured the distance between each fragment and the nearest settlement, continuous forest, and nearest neighboring fragment. We fit 10 candidate SAR models to the data using nonlinear least squares regression and compared them using Akaike's Information Criterion (AIC). To determine how habitat characteristics, as well as area, influenced species richness, we ran a hierarchical partitioning procedure to select which variables to include in generalized additive models (GAMs) and compared them using AIC. Contrary to expectations, we found that lemurs form convex SARs, without a "small island effect", and with the power model being the most likely SAR model. Although we found that four variables (area, survey effort, and total human disturbance, and mean tree height) independently contributed greater than 10% of the variation in lemur species richness, only area was included in the most likely model. We suggest that the power model was the most likely SAR model and our inability to detect a "small island effect" are the result of Microcebus spp. being edge tolerant and capable of dispersing through matrix, scale issues in the study design, and low γ-diversity in the landscape. However, more study is needed to determine what role human disturbance plays in influencing species richness in lemurs.

Fruit ripeness can be indicated through changes in chromaticity, luminance, odor, hardness, and size to attract seed dispersing animals. We quantified these attributes for both ripe and unripe fruits of 31 lemur-dispersed plant species in Ankarafantsika National Park, a tropical dry forest in northwestern Madagascar. We used spectroscopy, gas-chromatography mass-spectrometry, and a modified force gauge to quantify chromaticity, luminance, odor, and hardness. We compared these traits between unripe and ripe fruits of each species to determine which traits reliably indicate fruit ripeness across species. Overall, ripe fruits were significantly heavier and softer than unripe fruits. Ripe fruits were not more chromatically-conspicuous or odiferous relative to unripe fruits, nor were ripe fruits more conspicuous in the luminance channel. Contrary to expectation, our findings indicate that, in this particular system, plant-lemur interactions may be strongly mediated by haptic traits, such as fruit hardness, which are consistent and reliable indicators of fruit ripeness. Despite the potential importance of haptic indicators of fruit ripeness, they are underrepresented in the literature on sensory ecology.

Lactating females face energetic constraints absent in conspecifics and must compensate for higher energy requirements. Infant transport is the most energetically costly mammalian activity after lactation. Nonetheless, infant transport and mother-infant contact are seldom measured. The extreme seasonality characteristic of Madagascar coupled with lactation costs and infant transport is a trifold energetic challenge encountered by lemur mothers. We hypothesized that Coquerel's sifaka (Propithecus coquereli) mothers spend more time caring for infants during the early/earlier-mid lactation period, coinciding with the resource depletive austral winter, relative to later-mid/late lactation. We tested this hypothesis by measuring infant carrier identity, transport position, and frequency of mother-infant contact for 678 focal hours over two consecutive birth seasons from 1 to 26 weeks postnatal (N = 10 infants, N = 10 mothers, N = 19 adult males, N = 8 adult females) in Ankarafantsika National Park (ANP), Madagascar. Quantifying P. coquereli postnatal care strategies demonstrates how a species with a "slow" life history lives in an energetically challenging environment, thereby providing data on maternal energetic responses and infant development in an endangered strepsirrhine. Mothers were the primary infant transporters. Adult males and non-lactating females participated in infant transport, but for significantly less time. Infants spent significantly more time in the ventral transport position than dorsally or independently. Infants were still transported 26% of the time at 6.5 months postnatal. Infants initiated and broke bodily contact with mothers more frequently than mothers initiated and broke contact with infants. Infants were dependent on their mothers for longer durations than suggested by previous studies and carried dorsally until later ages than in other comparably sized wild lemurs. Am. J. Primatol. 78:646-658, 2016. © 2016 Wiley Periodicals, Inc.

Summary In contrast to the majority of primates, which exhibit dedicated diurnality or nocturnality, all species of Eulemur are cathemeral. Colour vision, in particular, is strongly affected by the spectral composition and intensity of ambient light, and the impact of activity period on the evolution of primate colour vision is actively debated. We studied three groups of wild brown lemurs (Eulemur fulvus) in Ankarafantsika National Park, Madagascar, over a 1-year span. We also used non-invasive faecal DNA collection and analysis to study the opsin genes underlying the colour vision of 24 individuals. We quantified the colour and brightness of dietary fruits and modelled the chromaticity and discriminability of food objects to different visual phenotypes under daylight, twilight and moonlight conditions. We found that E. fulvus possesses routine dichromacy, unlike its congener E. flavifrons, for which polymorphic trichromacy has been reported. Our models suggest that dichromacy is well suited to the foraging ecology of E. fulvus. The performance of modelled dichromats and trichromats is comparable under nocturnal illuminants, and the luminance values of most diet items are detectable across light conditions. The trichromatic phenotype demonstrates a modest advantage under daylight conditions. Our results, taken together with reports of polymorphic trichromacy in E. flavifrons, suggest functional ecological variation in the visual system of the genus Eulemur. Interspecific phenotypic variation in the colour vision of a genus is both unexpected and instructive. Ecological differences between species of Eulemur could reveal thresholds for the origins of polymorphic trichromacy, which preceded the evolution of routine trichromatic vision in humans and other primates.

Animal reliance on fruit signals, such as hardness, colour, and odour, during foraging is poorly understood. Here, we present data on fruit foraging behaviour and efficiency (rate of fruit ingestion) of three groups of wild, frugivorous brown lemurs (Eulemur fulvus, N = 29 individuals) in Ankarafantsika National Park, Madagascar. We quantify fruit hardness using a modified force gauge, fruit colour using spectroscopy, and fruit odour using volatile organic compound (VOC) sampling with gas chromatography-mass spectrometry. We relate lemur foraging behaviour to fruit traits by calculating touching, visual inspection, and sniffing indices and relate lemur foraging efficiency to fruit traits by calculating acceptance indices. The use of different sensory modalities by lemurs is marginally predicted in one case by fruit traits—fruits with higher overall smell signals are sniffed less than fruits with lower overall smell signals. When controlling for all fruit traits, fruit size is the only significant predictor of fruit foraging efficiency—lemurs forage more rapidly on smaller fruits relative to larger fruits.

Rapid deforestation has fragmented habitat across the landscape of Madagascar. To determine the effect of fragmentation on seed banks and the potential for forest regeneration, we sampled seed viability, density and diversity in 40 plots of 1 m2 in three habitat types: forest fragments, the near edge of continuous forest, and deforested savanna in a highly fragmented dry deciduous forest landscape in northwestern Madagascar. While seed species diversity was not different between forest fragments and continuous forest edge, the number of animal-dispersed seeds was significantly higher in forest fragments than in continuous forest edge, and this pattern was driven by a single, small-seeded species. In the savanna, seeds were absent from all but three of the 40 plots, indicating that regeneration potential is low in these areas. Several pre- and post-dispersal biotic and abiotic factors, including variation in the seed predator communities and edge effects could explain these findings. Understanding the extent to which seed dispersal and seed banks influence the regeneration potential of fragmented landscapes is critical as these fragments are the potential sources of forest expansion and re-connectivity. Resume Le sort de graines dispersees peut etre altere en reponse a la perte et la fragmentation d’ habitats, modifiant ainsi les banques de semences du sol. Nous avons echantillonne la viabilite, la densite et la diversite des graines dans 40 parcelles de 1 m2 en trois types d'habitats dans le paysage forestier feuillu, sec, et fragmente du nord-ouest de Madagascar: (1) fragments de foret (2) bords rapproches de foret ininterrompue, et (3) savane deboisee. Notre objectif etait d’etablir l'effet de cette fragmentation sur les banques de semences, et la possibilite de regeneration forestier. Nos resultats demontrent, bien que la diversite des especes de semences ne varie pas entre les fragments de foret et les bords rapproches de foret ininterrompue, que la quantite de graines dispersees par les animaux est considerablement plus elevee dans le premier type d'habitat que dans la deuxieme, et que cet effet est grâce a une seule espece [a petites graines]. Dans la savane deboisee, une absence complete de graines a ete notee dans tous les 40 parcelles, a part trois. Plusieurs facteurs biotiques ainsi qu’ abiotiques, pre- et post-dispersion, sont impliques, y compris des variations au sein des communautes de predateurs de graines ainsi que des effets de lisiere. Le potentiel de regeneration en paysages fragmentes repose sur une meilleure comprehension de la nature et l’etendue des changements dans la dispersion des graines, ainsi que les resultantes banques de semences. Ceci est particulierement important au Madagascar, ou la perte des forets se poursuit a un rythme precipite, puisque ces fragments sont les sources essentielles de l'expansion et de la reconnexion forestiere.

Pitheciids are known for their frugivorous diets, but there has been no broad-scale comparison of fruit genera used by these primates that range across five geographic regions in South America. We compiled 31 fruit lists from data collected from 18 species (three Cacajao, six Callicebus, five Chiropotes, and four Pithecia) at 26 study sites in six countries. Together, these lists contained 455 plant genera from 96 families. We predicted that 1) closely related Chiropotes and Cacajao would demonstrate the greatest similarity in fruit lists; 2) pitheciids living in closer geographic proximity would have greater similarities in fruit lists; and 3) fruit genus richness would be lower in lists from forest fragments than continuous forests. Fruit genus richness was greatest for the composite Chiropotes list, even though Pithecia had the greatest overall sampling effort. We also found that the Callicebus composite fruit list had lower similarity scores in comparison with the composite food lists of the other three genera (both within and between geographic areas). Chiropotes and Pithecia showed strongest similarities in fruit lists, followed by sister taxa Chiropotes and Cacajao. Overall, pitheciids in closer proximity had more similarities in their fruit list, and this pattern was evident in the fruit lists for both Callicebus and Chiropotes. There was no difference in the number of fruit genera used by pitheciids in habitat fragments and continuous forest. Our findings demonstrate that pitheciids use a variety of fruit genera, but phylogenetic and geographic patterns in fruit use are not consistent across all pitheciid genera. This study represents the most extensive examination of pitheciid fruit consumption to date, but future research is needed to investigate the extent to which the trends in fruit genus richness noted here are attributable to habitat differences among study sites, differences in feeding ecology, or a combination of both.

Edge effects are an inevitable and important consequence of forest loss and fragmentation. These effects include changes in species biology and biogeography. Here we examine variations in body mass and morphometrics for 2 sympatric species of mouse lemurs (Microcebus murinus and M. ravelobensis) between edge and interior habitats in the dry deciduous forest at Ankarafantsika National Park. Between May and August 2012, we conducted mark-recapture experiments on mouse lemurs trapped along edge and interior forest transects within continuous forest adjacent to a large savannah. Of the 34 M. murinus captured during our study, 82% (n = 28) were trapped in interior habitats. Conversely, 72% (n = 47) of M. ravelobensis were captured in edge habitats. We found that mean body mass of M. murinus and M. ravelobensis did not differ between edge and interior habitats. However, female M. ravelobensis weighed significantly more in edge habitats (56.09 ± 1.74 g) than in interior habitats (48.14 ± 4.44 g). Our study provides some of the first evidence of sex differences in edge responses for a primate species.

Introduction - Malagasy habitats undergo vast seasonal changes in temperature and rainfall regimes, which in turn strongly infl uence the biology and life history of their inhabitants (Wright, 1999 ). The western dry deciduous forests show a yearly alternation between a distinct dry season and a hot and humid rainy season (Jury, 2003 ). Correspondingly, the phenology of dry forest trees – i.e., the availability of leaves, fl owers and fruits – changes substantially over the course of a year (Sorg and Rohner, 1996 ; Thoren et al ., 2011 ). As a result, plant food resources are not available evenly throughout the yearly cycle, and a relatively resource-poor dry season can be distinguished from a relatively resource-rich rainy season (Hladik et al ., 1980 ). The length of both seasons varies across Madagascar due to a geographical gradient in minimal temperatures and annual rainfall . Adaptive responses of lemurs to this environmental seasonality include seasonal changes in general activity (Schmid and Kappeler, 1998 ; Wright, 1999 ; Dausmann et al ., 2004 ), shelter usage pattern (Ramanankirahina et al ., 2012 ), body mass (Hladik et al ., 1980 ; Schmid and Kappeler, 1998 ; Atsalis, 1999 ; Randrianambinina et al ., 2003 ), reproduction (Hladik et al ., 1980 ; Wright, 1999 ), and feeding regime (Hladik et al ., 1980 ; Thoren et al ., 2011 ). It is known, for example, that diademed sifakas ( Propithecus diadema ) shift their diet from fruits and seeds to non-fruit foods, such as leaves and fl owers, and decrease food ingestion and general feeding time during the dry season (Irwin et al ., 2014 ). This ability to fl exibly change behavior or resource usage patterns in view of certain environmental changes is the result of adaptive plasticity (Nussey et al ., 2007 ). Adaptive plasticity in seasonal feeding regimes can be expected to be highest in omnivorous species that can, in principle, switch between different sources of energy and protein by consuming different food plant species or varying the proportion of animal prey in the diet. Moreover, omnivores can consume different parts, such as fruits and gum s, of the same food plant species according to the availability of these items.

The dwarf and mouse lemurs of Madagascar are two very species-rich lemur genera, yet there is a relative paucity of information on this primate family in published literature. In this first ever treatment of the Cheirogaleidae, international experts are brought together to review and integrate our current knowledge of the behaviour, physiology, ecology, genetics and biogeography of these species. A wide range of direct and indirect research methods that are currently used to study these cryptic nocturnal solitary foragers are described. By uniting often disparate research on captive and free-ranging taxa and synthesising recent methodological advances, this book provides new insights that will encourage further studies of this fascinating primate family. This synthesis will provide an incentive for more integrative studies of the Cheirogaleidae in captivity and in the wild, enabling the impacts of deforestation and other factors to be identified and directions for future conservation efforts to be established.

Neotropical monkeys of the genera Cacajao, Chiropotes, and Pithecia (Pitheciidae) are considered to be highly arboreal, spending most of their time feeding and traveling in the upper canopy. Until now, the use of terrestrial substrates has not been analyzed in detail in this group. Here, we review the frequency of terrestrial use among pitheciin taxa to determine the ecological and social conditions that might lead to such behavior. We collated published and unpublished data from 14 taxa in the three genera. Data were gleaned from 53 published studies (including five on multiple pitheciin genera) and personal communications of unpublished data distributed across 31 localities. Terrestrial activity was reported in 61% of Pithecia field studies (11 of 18), in 34% of Chiropotes studies (10 of 29), and 36% of Cacajao studies (4 of 11). Within Pithecia, terrestrial behavior was more frequently reported in smaller species (e.g. P. pithecia) that are vertical clingers and leapers and make extensive use of the understory than in in the larger bodied canopy dwellers of the western Amazon (e.g. P. irrorata). Terrestrial behavior in Pithecia also occurred more frequently and lasted longer than in Cacajao or Chiropotes. An apparent association was found between flooded habitats and terrestrial activity and there is evidence of the development of a “local pattern” of terrestrial use in some populations. Seasonal fruit availability also may stimulate terrestrial behavior. Individuals also descended to the ground when visiting mineral licks, escaping predators, and responding to accidents such as a dropped infant. Overall, the results of this review emphasize that terrestrial use is rare among the pitheciins in general and is usually associated with the exploitation of specific resources or habitat types. Am. J. Primatol. 74:1106-1127, 2012. © 2012 Wiley Periodicals, Inc.

We tested four major hypotheses on the ecological aspects of body mass variation in extant Mala- gasy strepsirrhines: thermoregulation, resource seasonal- ity/scarcity, resource quality, and primary productivity. These biogeographic hypotheses focus on the ecological aspects of body mass variation, largely ignoring the role of phylogeny for explaining body mass variation within line- ages. We tested the independent effects of climate and resource-related variables on variation in body mass among Malagasy primates using recently developed com- parative methods that account for phylogenetic history and spatial autocorrelation. We extracted data on lemur body mass and climate variables for a total of 43 species from 39 sites. Climatic data were obtained from the WorldClim database, which is based on climate data from weather stations compiled around the world. Using gener- alized linear models that incorporate parameters to account for phylogenetic and spatial autocorrelation, we found that diet and climate variables were weak predic- tors of lemur body mass. Moreover, there was a strong phylogenetic effect relative to the effects of space on lemur body mass in all models. Thus, we failed to find support for any of the four hypotheses on patterns of geography and body mass in extant strepsirrhines. Our results indi- cate that body mass has been conserved since early in the evolutionary history of each genus, while species diversi- fied into different environmental niches. Our findings are in contrast to some previous studies that have suggested resource and climate related effects on body mass, though these studies have examined this question at different taxonomic and/or geographic scales. Am J Phys Anthropol 147:401-408, 2012. V

Prolemur simus (the greater bamboo lemur) is the most abundant lemur in the northern subfossil sites of Madagascar. Living populations still persist, but in low numbers within a diminished range, making it one of the most critically endangered lemurs. Over the past twenty years scientists have searched the south- and central-eastern rain forests of Madagascar. Despite surveys that encompass over 500 km2, less than 75 animals have been found, with a recent total count of 60. More encouraging is that in 2007 two new sites containing P. simus were found: Mahasoa an unprotected 150 ha fragment east of the Ranomafana/Andringitra corridor (17 P. simus), and Torotorofotsy, a RAMSAR site near Andasibe (∼16 P. simus). Prolemur simus is a bamboo specialist with a patchy geographic distribution, which may be driven by the distribution of one or two bamboo species. Home ranges are large, group size has been observed to be from four to 26 individuals, and localities may be spaced hundreds of kilometers apart...

We determined if data on strepsirhine body and home range sizes support an optimal body size (OBS) model of 100 g, as predicted from studies of energetics in terrestrial mammals. We also tested the following predictions of the OBS model: 1) relationships between body and home range sizes will change slope and sign above and below the OBS threshold of 100 g and 2) best-fit lines for OBS regression models (above and below the 100-g threshold) will intersect at ca. 100 g (range of 80–250 g). We collected data on body mass, home range size, and vertical ranging behavior for 37 strepsirhines from the literature. Linear regression analyses and phylogenetic independent contrasts methods revealed that body size is a significant determinant of both 2-dimensional (ha) and 3-dimensional (km3) home range sizes only in taxa weighing >100 g. There were consistent changes in the sign of the slopes above and below the OBS threshold. The intersections of the best-fit lines were within the OBS range for the body size to 3-dimensional home range comparisons. Thus, the data provide some support for the OBS model in strepsirhines. However, no regression model was statistically significant for the taxa below the OBS threshold, which may reflect small sample sizes. Also, no slope differed significantly between taxa above and below the OBS. Significant correlations between body and home range sizes for the complete data sets refute the √-shaped constraint space predicted via the OBS model.

I investigated ecological and phylogenetic correlates to body size variations in 10 taxa of extant Indriidae (Indri, Avahi, and Propithecus). I also tested for phylogenetic niche conservatism as a model for the evolution of indriid body size. Phylogenetic niche conservatism refers to the shared attributes that related taxa have acquired because they tend to have occupied similar niches during their evolutionary history. I collected species-specific data on body mass, climate, density, and chemical properties of food items from the literature. I used 2 phylogenies in independent contrasts methods to control for phylogenetic relationships (Indri and Propithecus as sister taxa vs. Indri basal taxa to all indriids). Multivariate models indicated that lemur density and resource quality are the strongest ecological correlates to indriid body size variations. Partitioning methods revealed that 52.4–67% of indriid body size variation is explained by phylogenetic niche conservation. Thus, indriid body size variations may be the result of stabilizing selection. Though it is possible to identify constraints on lower than average body size, there are few data on selection against larger than average body size in indriids. Large body size in subfossil lemurs further complicates identification of constraints on larger than average body size in extant indriids. Researchers using independent contrast methods to control for phylogeny should be aware that some ecology-phenotype relationships are best explained as the result of the synergistic effects of ecology and phylogeny.

Lactating females face energetic constraints absent in conspecifics and must compensate for higher energy requirements. Infant transport is the most energetically costly mammalian activity after lactation. Nonetheless, infant transport and mother-infant contact are seldom measured. The extreme seasonality characteristic of Madagascar coupled with lactation costs and infant transport is a trifold energetic challenge encountered by lemur mothers. We hypothesized that Coquerel's sifaka (Propithecus coquereli) mothers spend more time caring for infants during the early/earlier-mid lactation period, coinciding with the resource depletive austral winter, relative to later-mid/late lactation. We tested this hypothesis by measuring infant carrier identity, transport position, and frequency of mother-infant contact for 678 focal hours over two consecutive birth seasons from 1 to 26 weeks postnatal (N = 10 infants, N = 10 mothers, N = 19 adult males, N = 8 adult females) in Ankarafantsika National Park (ANP), Madagascar. Quantifying P. coquereli postnatal care strategies demonstrates how a species with a "slow" life history lives in an energetically challenging environment, thereby providing data on maternal energetic responses and infant development in an endangered strepsirrhine. Mothers were the primary infant transporters. Adult males and non-lactating females participated in infant transport, but for significantly less time. Infants spent significantly more time in the ventral transport position than dorsally or independently. Infants were still transported 26% of the time at 6.5 months postnatal. Infants initiated and broke bodily contact with mothers more frequently than mothers initiated and broke contact with infants. Infants were dependent on their mothers for longer durations than suggested by previous studies and carried dorsally until later ages than in other comparably sized wild lemurs. Am. J. Primatol. 78:646-658, 2016. © 2016 Wiley Periodicals, Inc.

We investigated how greater dwarf lemur (Cheirogaleus major) densities, ambient air temperature, and dendrometrics (tree height and diameter) varied along forest edge-interior gradients in the Vohibola III Classified Forest in SE Madagascar. We also assessed if spatial variations in densities of Cheirogaleus major provide indirect evidence of increased predation pressure in the transition zone between edge and interior forest habitats, i.e., an ecological trap. We conducted diurnal temperature surveys (N = 394) and nocturnal surveys of Cheirogaleus major (N = 182) over 2 yr along 4 1250-m transects that ran perpendicular to the forest edge in Vohibola III. We did not see Cheirogaleus major from May to mid-September, and the highest sighting frequency occurred during October–November. Cheirogaleus major exhibited a negative edge response because densities ranged from low levels in edge habitats to higher levels in the forest interior. After we tested for spatial autocorrelation, edge-related variations in densities of Cheirogaleus major covaried most strongly with tree diameter. Edge responses of Cheirogaleus major may reflect spatial variations in fruit and liana abundance, though data are needed on the precise relationship between tree diameter and food production to confirm the relationship. Edge-related variations in densities of Cheirogaleus major may also provide indirect evidence of an ecological trap. Testing and controlling for spatial autocorrelation should be important components of future studies of primate conservation biology and ecology.

I investigated how transect type (trails vs. cut transects) and seasonality influenced density estimates for 5 lemur taxa (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus) in the Vohibola III Classified Forest in SE Madagascar. I surveyed tree height and diameter and lemur populations from June 1 to December 28, 2004 along 2 1250-m trails local people used and 2 1250-m transects cut parallel to the trails in primary rain forest. Despite dendrometric variations within and between trails and transects, only density estimates of Hapalemur griseus griseus differed significantly by transect type. The spatial variation may be a result of removal by local people of giant bamboo, which is the main food for Hapalemur griseus griseus, along trails. Conversely, seasonality influenced density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus. The temporal variations may be related to seasonal torpor for Cheirogaleus major and increased detection probabilities during periods of fruit exploitation for Eulemur rubriventer, Microcebus rufus, and Hapalemur griseus griseus. Transect type and seasonality did not affect density estimates for Avahi laniger, which may be related to the highly folivorous and low-energy diet of the nocturnal lemur. Researchers surveying lemurs along line transects should be aware that transect selection may influence density estimates for Hapalemur griseus griseus and that seasonality may influence density estimates for Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, and Microcebus rufus.

The Fandriana-Marolambo forest corridor is one of the largest (ca. 250,000 ha) and least explored tracts of unprotected forest in southeast Madagascar. Although published range maps show continuous distributions for many lemurs throughout the region, there are few data on lemur community structure in the corridor. We aimed to determine lemur community structure, with its ecological correlates (altitude, agriculture, selective logging, and hunting), in the Fandriana-Marolambo forest corridor. We surveyed 7 sites and sighted 4 nocturnal taxa (Avahi laniger, Cheirogaleus major, Lepilemur mustelinus, and Microcebus rufus) and 6 diurnal taxa (Eulemur rubriventer, E. fulvus rufus, E. f. fulvus, Propithecus diadema edwardsi, Hapalemur griseus griseus, and Varecia variegata variegata). Composition of the lemur community was broadly similar to that of nearby protected areas (Ranomafana and Mantadia National Parks). However, we sighted no Hapalemur aureus, H. simus, or Indri indri, and observed Propithecus diadema edwardsi and Varecia variegata variegata at only 1 site each. We sighted an apparent hybrid form of Eulemur fulvus fulvus and E. f. rufus that may represent a new hybrid zone for lemurs. After testing for spatial autocorrelation, lemur diversity correlates negatively with altitude and agricultural intensity. Though the Government of Madagascar is assessing the corridor as a new national park, we suggest conservation plans for local lemurs are complicated by population isolation and lack of data on minimum viable size of the proposed protected area.

I present data on variations in Eulemur fulvus rufus and Lepilemur mustelinus densities as well as tree characteristics (height, diameter and stem frequency) between edge and interior forest habitats in southeastern Madagascar. Line transect surveys were conducted from June 2003 to November 2005 in edge and interior forest habitats in the Vohibola III Classified Forest. Although E. f. rufus densities were significantly lower in edge habitats than in interior habitats, density estimates for L. mustelinus did not differ significantly between habitats. Trees in edge habitats were significantly shorter, had smaller diameters and had lower stem frequencies (for those 25 cm in diameter) than trees in interior habitats. Spatial characteristics of food abundance and quality may explain lemur density patterns in Vohibola III. Low E. f. rufus densities may reduce seed dispersal in edge habitats, which has important consequences for the long-term viability of forest ecosystems in Madagascar.

Edge effects are caused by the penetration of abiotic and biotic conditions from the matrix into forest interiors. Although edge effects influence the biogeography of many tropical organisms, they have not been studied directly in primates. Edge effects are particularly relevant to lemurs due to the loss of 80-90% of forests in Madagascar. In this study, data are presented on how biotic edge effects influenced the distribution and density of lemurs in the Vohibola III Classified Forest in southeastern Madagascar. In total, 415 lemur surveys were conducted during June-October 2003 and May-September 2004 along six 1,250-m transects that ran perpendicular to the forest edge. Data were also collected on lemur food trees along the six transects (density, height, diameter at breast height, area, volume, and distance to forest edge). Four nocturnal species (Avahi laniger, Cheirogaleus major, Lepilemur microdon, and Microcebus rufus) and four diurnal species (Eulemur rubriventer, Eulemur fulvus rufus, Hapalemur grisesus griseus, and Propithecus diadema edwardsi) were sighted during surveys. Regression analyses of lemur densities as a function of distance to forest edge provided edge tolerances for A. laniger (edge-tolerant), M. rufus (edge-tolerant), E. rubriventer (edge-tolerant or omnipresent), and H. g. griseus (omnipresent). The density and distribution of M. rufus and their foods trees were correlated. Edge-related variations in food quality and predation pressures may also be influencing lemurs in Vohibola III. Tolerance for edge effects may explain, in part, how lemurs have survived extreme habitat loss and forest fragmentation in southeastern Madagascar.

This paper describes the results of field surveys for two Endangered lemurs, Milne-Edward's sifaka Propithecus diadema edwardsi and the black and white ruffed lemur Varecia variegata variegata, in the unprotected Fandriana-Marolambo forest corridor in south-east Madagascar. Published range maps show P. d. edwardsi and V. v. variegata present in the corridor, although few surveys have been conducted in this area. A total of 292.9 km of surveys were conducted at eight sites in the corridor but P. d. edwardsi and V. v. variegata were each located at only one site. Absence of these species at other sites may be the combined result of altitudinal and anthropogenic effects. Conservation plans for P. d. edwardsi and V. v. variegata are complicated because gaps in forest cover prevent dispersal of conspecifics from nearby protected areas. Conservation authorities should be cautious in using range maps based on the presumed distribution of lemur species.

Species that exploit a wide range of resources or habitats (generalists) tend to be widely distributed, whereas species that exploit a narrow range of resources or habitats (specialists) often have a limited distribution. The distribution patterns are thought to result from specialists using relatively smaller habitats than those exploited by generalists. I used data from 1,725 km of primate surveys that I conducted in Guyana to test these hypotheses. Habitat breadth is the total number of different habitat types occupied by each species. I used the total number of different food categories exploited by each species to measure dietary breadth. Geographic range size is correlated with habitat breadth but not with dietary breadth or body size for the 8 primate species in Guyana. Habitat generalists—red howlers and wedge-capped capuchins—range into all habitats. Habitat specialists—spider monkeys, brown bearded sakis, and golden-handed tamarins—range only into large habitats. Habitat generalists tend to be dietary type specialists in Guyana. I suggest that only habitat generalists can subsist on the low-quality foods in small habitats in Guyana. Conversely, habitat specialists tend to be dietary type generalists in Guyana. They must feed on a variety of food types in large habitats. However, using the number of food categories exploited as a measure of dietary breadth may be only a weak aspect of multidimensional niche. Researchers testing biogeographic hypotheses associated with dietary breadth should consider including multivariate indicators of both the types of food categories eaten and the number of plant species exploited.

Species-area relationships predict that there is a positive relationship between the number of species and the size of an area. It has been suggested that species richness will covary with area because larger areas have a greater diversity of habitats. Moreover, habitat diversity may operate in conjunction with riverine barriers to influence primate biogeography. Few studies have determined if and how these hypotheses relate to primate diversity in Guyana. To test these biogeographic hypotheses, I used data from 1,725 km of primate surveys I conducted in Guyana. I estimated geographic ranges for each of the 8 primate species via a GIS system. Geographic range size is a major determinant of the number of sightings of the 8 primate species. Primate species diversity is strongly negatively correlated with the number of rivers crossed moving in a clockwise pattern from eastern to NW Guyana. Interfluvial and habitat areas influence primate species diversity in Guyana. However, my data on primate biogeography in Guyana do not support the hypothesis that habitat diversity within the interfluvial areas effects primate diversity. Although the species-area relationship is considered the closest thing to a rule in ecology, researchers should be wary of too readily applying and accepting the model at all scales in biogeographic studies.

The island of Madagascar is characterized by high levels of biodiversity and endemism, with at least 70 % of the 12,000 plant species and all of the approximately 93 primate species found nowhere else in the world. These biogeographic patterns are remarkable given that on-going climate change and anthropogenic disturbances have resulted in the conversion of 40 % of the original forest cover into grasslands and agricultural fields. This incongruence between high lemur diversity and anthropogenic disturbance provides for an ideal natural setting for research on the effects of altitude on lemur conservation biogeography. In addition to reviewing lemur biogeography as it pertains to altitude, I investigated the biogeography of lemur species diversity and assemblages at 16 sites in montane and cloud forests in southeast Madagascar. Although lemur species diversity showed a significant negative relationship with site elevation, this relationship held only for sites lower than 1,500 m elevation. There was strong evidence for nestedness in terms of the effects of altitudinal variation on lemur assemblage patterns across the sites, indicating that those species that range into the relatively high-altitude sites are a subset of those that range in relatively low altitude sites. This nestedness is likely due to the combined effects of lower temperatures and reduced resource abundance at higher elevations. Lemurs face an uncertain future across the elevation gradients within this landscape, irrespective of their tolerance for high-altitude habitats.