Effects of Forest Fragmentation on Connectivity and Genetic Diversity in an Endemic and an Invasive Rodent in Northwestern Madagascar.

Habitat loss and fragmentation are of concern to conservation biologists worldwide. However, not all organisms are affected equally by these processes; thus, it is important to study the effects of living in fragmented habitats on species that differ in lifestyle and habitat requirements. In this study, we examined the dispersal and connectivity patterns of rodents, one endemic (Eliurus myoxinus) and one invasive (Rattus rattus), in two landscapes containing forest fragments and adjacent continuous forest patches in northwestern Madagascar. We generated genetic (RADseq) data for 66 E. myoxinus and 81 R. rattus individuals to evaluate differences in genetic diversity as well as inbreeding and connectivity in two landscapes. We found higher levels of inbreeding and lower levels of genetic diversity in E. myoxinus compared with R. rattus. We observed related dyads both within and between habitat patches and positive spatial autocorrelation at lower distance classes for both species, with a stronger pattern of spatial autocorrelation in R. rattus. Across each site, we identified contrasting migration rates for each species, but these did not correspond to habitat-matrix dichotomies. The relatively low genetic diversity in the endemic E. myoxinus suggests ecological constraints that require further investigation.

The burning island: Spatiotemporal patterns of fire occurrence in Madagascar.

Anthropogenic fire use is widespread across Madagascar and threatens the island's unprecedented endemic biodiversity. The vast majority (96%) of lemur species are already threatened with extinction, and Madagascar has already lost more than 44% of its forests. Previous conservation assessments have noted the role of fire in the rampant deforestation and habitat degradation across Madagascar, but published, quantified data on fire use across the island are incredibly limited. Here, we present the first quantification of spatiotemporal patterns in fire occurrence across Madagascar using VIIRS satellite fire detection data. We assess which regions of Madagascar have the most prevalent fire use, how fire use is changing over time, and what this means for Madagascar's remaining forest ecosystems. An average of 356,189 fires were detected every year in Madagascar from 2012-2019, averaging 0.604 fires/km2. Fire use was near-ubiquitous across the island, but was most prevalent in the western dry deciduous forests and succulent woodlands ecoregions. Fire frequency in the eastern lowlands was highest around the remaining humid rainforest, and fire frequency was increasing over time around much of the remaining humid and dry forest. We found that 18.6% of all remaining forest was within 500 m of a fire within a single year, and 39.3% was within 1 km. More than half of remaining forest was within 1 km of a fire in a single year in the dry deciduous forests, succulent woodlands, and mangroves ecoregions. However, fire frequency within national park protected areas was, on average, 65% lower than their surroundings. Only 7.1% of national park forest was within 500 m of a fire within one year, and 17.1% was within 1 km, suggesting that national parks are effective at reducing fire frequency in Madagascar's tropical forests.

Enter the Matrix: Use of Secondary Matrix by Mouse Lemurs.

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.

Effects of habitat edges on vegetation structure and the vulnerable golden-brown mouse lemur (Microcebus ravelobensis) in northwestern Madagascar.

BACKGROUND: Edge effects can influence species composition and community structure as a result of changes in microenvironment and edaphic variables. We investigated effects of habitat edges on vegetation structure, abundance and body mass of one vulnerable Microcebus species in northwestern Madagascar. We trapped mouse lemurs along four 1000-m transects (total of 2424 trap nights) that ran perpendicular to the forest edge. We installed 16 pairs of 20 m2 vegetation plots along each transect and measured nine vegetation parameters. To determine the responses of the vegetation and animals to an increasing distance to the edge, we tested the fit of four alternative mathematical functions (linear, power, logistic and unimodal) to the data and derived the depth of edge influence (DEI) for all parameters. RESULTS: Logistic and unimodal functions best explained edge responses of vegetation parameters, and the logistic function performed best for abundance and body mass of M. ravelobensis. The DEI varied between 50 m (no. of seedlings, no. of liana, dbh of large trees [dbh ≥ 10 cm]) and 460 m (tree height of large trees) for the vegetation parameters, whereas it was 340 m for M. ravelobensis abundance and 390 m for body mass, corresponding best to the DEI of small tree [dbh < 10 cm] density (360 m). Small trees were significantly taller and the density of seedlings was higher in the interior than in the edge habitat. However, there was no significant difference in M. ravelobensis abundance and body mass between interior and edge habitats, suggesting that M. ravelobensis did not show a strong edge response in the study region. Finally, regression analyses revealed three negative (species abundance and three vegetation parameters) and two positive relationships (body mass and two vegetation parameters), suggesting an impact of vegetation structure on M. ravelobensis which may be partly independent of edge effects. CONCLUSIONS: A comparison of our results with previous findings reveals that edge effects are variable in space in a small nocturnal primate from Madagascar. Such an ecological plasticity could be extremely relevant for mitigating species responses to habitat loss and anthropogenic disturbances.

Occurrence of lemurs in landscapes and their species-specific scale responses to habitat loss.

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.

Ecological fragmentation effects in mouse lemurs and small mammals in northwestern Madagascar.

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.

Species-area relationships of lemurs in a fragmented landscape in Madagascar.

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.

Lemur species-specific metapopulation responses to habitat loss and fragmentation.

Determining what factors affect species occurrence is vital to the study of primate biogeography. We investigated the metapopulation dynamics of a lemur community consisting of eight species (Avahi occidentalis, Propithecus coquereli, Microcebus murinus, Microcebus ravelobensis, Lepilemur edwardsi, Cheirogaleus medius, Eulemur mongoz, and Eulemur fulvus) within fragmented tropical dry deciduous forest habitat in Ankarafantsika National Park, Madagascar. We measured fragment size and isolation of 42 fragments of forest ranging in size from 0.23 to 117.7 ha adjacent to continuous forest. Between June and November 2011, we conducted 1218 surveys and observed six of eight lemur species (M. murinus, M. ravelobensis, C. medius, E. fulvus, P. coquereli, and L. edwardsi) in the 42 fragments. We applied among patch incidence function models (IFMs) with various measures of dispersal and a mainland-island IFM to lemur species occurrence, with the aim of answering the following questions: 1) Do lemur species in dry deciduous forest fragments form metapopulations? 2) What are the separate effects of area (extinction risk) and connectivity/isolation (colonization potential) within a lemur metapopulation? 3) Within simulated metapopulations over time, how do area and connectivity/isolation affect occurrence? and 4) What are the conservation implications of our findings? We found that M. murinus formed either a mainland-island or an among patch metapopulation, M. ravelobensis formed a mainland-island metapopulation, C. medius and E. fulvus formed among patch metapopulations, and neither P. coquereli or L. edwardsi formed a metapopulation. Metapopulation dynamics and simulations suggest that area was a more consistent positive factor determining lemur species occurrence than fragment isolation and is crucial to the maintenance of lemur populations within this fragmented landscape. Using a metapopulation approach to lemur biogeography is critical for understanding how lemur species respond to forest loss and fragmentation.

Infant transport and mother-infant contact from 1 to 26 weeks postnatal in Coquerel's sifaka (Propithecus coquereli) in northwestern Madagascar.

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.

Edge effects on morphometrics and body mass in two sympatric species of mouse lemurs in Madagascar.

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.

Colour and odour drive fruit selection and seed dispersal by mouse lemurs.

Animals and fruiting plants are involved in a complex set of interactions, with animals relying on fruiting trees as food resources, and fruiting trees relying on animals for seed dispersal. This interdependence shapes fruit signals such as colour and odour, to increase fruit detectability, and animal sensory systems, such as colour vision and olfaction to facilitate food identification and selection. Despite the ecological and evolutionary importance of plant-animal interactions for shaping animal sensory adaptations and plant characteristics, the details of the relationship are poorly understood. Here we examine the role of fruit chromaticity, luminance and odour on seed dispersal by mouse lemurs. We show that both fruit colour and odour significantly predict fruit consumption and seed dispersal by Microcebus ravelobensis and M. murinus. Our study is the first to quantify and examine the role of bimodal fruit signals on seed dispersal in light of the sensory abilities of the disperser.

Edge effects on foliar stable isotope values in a Madagascan tropical dry forest.

Edge effects represent an inevitable and important consequence of habitat loss and fragmentation. These effects include changes in microclimate, solar radiation, or temperature. Such abiotic effects can, in turn, impact biotic factors. They can have a substantial impact on species, communities, and ecosystems. Here we examine clinal variations in stable carbon and nitrogen isotope values for trees along an edge-interior gradient in the dry deciduous forest at Ankarafantsika National Park. We predicted that soil respiration and differences in solar irradiance would result in stratified δ¹³C values where leaves collected close to the forest floor would have lower δ¹³C values than those growing higher up in the canopy. We also anticipated that plants growing at the savannah-forest boundary would have higher δ¹³C and δ¹5N values than plants growing in the forest interior. As expected, we detected a small but significant canopy effect. Leaves growing below 2 m from the forest floor exhibit δ¹³C values that are, on average, 1.1‰ lower than those growing above this threshold. We did not, however, find any relationship between foliar δ¹³C and distance from the edge. Unpredictably, we detected a striking positive relationship between foliar δ¹5N values and increasing distance into the forest interior. Variability in physiology among species, anthropogenic influence, organic input, and rooting depth cannot adequately explain this trend. Instead, this unexpected relationship most likely reflects decreasing nutrient or water availability, or a shift in N-sources with increasing distance from the savannah. Unlike most forest communities, the trees at Ampijoroa are growing in nutrient-limited sands. In addition to being nutrient poor, these well-drained soils likely decrease the amount of soil water available to forest vegetation. Continued research on plant responses to edge effects will improve our understanding of the conservation biology of forest ecosystems in Madagascar.

Testing Bergmann's rule and the resource seasonality hypothesis in Malagasy primates using GIS-based climate data.

We tested four major hypotheses on the ecological aspects of body mass variation in extant Malagasy strepsirrhines: thermoregulation, resource seasonality/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 lineages. We tested the independent effects of climate and resource-related variables on variation in body mass among Malagasy primates using recently developed comparative 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 generalized linear models that incorporate parameters to account for phylogenetic and spatial autocorrelation, we found that diet and climate variables were weak predictors 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 indicate that body mass has been conserved since early in the evolutionary history of each genus, while species diversified 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.

Spatial variations in Eulemur fulvus rufus and Lepilemur mustelinus densities in Madagascar.

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.

Lemur responses to edge effects in the Vohibola III classified forest, Madagascar.

Forest edges are dynamic zones characterized by the penetration (to varying depths and intensities) of conditions from the surrounding environment (matrix) into the forest interior. Although edge effects influence many tropical organisms, they have not been studied directly in primates. Edge effects are particularly relevant to lemurs because of the highly fragmented forest landscapes found in Madagascar. In this study, data are presented regarding how the densities of six lemur species (Avahi laniger, Cheirogaleus major, Eulemur rubriventer, Hapalemur griseus griseus, Microcebus rufus, and Propithecus diadema edwardsi) varied between six 500-m interior transects and six 500-m edge transects in the Vohibola III Classified Forest in SE Madagascar. Diurnal (n = 433) and nocturnal (n = 128) lemur surveys were conducted during June-October 2003 and May-November 2004. A. laniger, E. rubriventer, and H. g. griseus exhibited a neutral edge response (no differences in densities between habitats). M. rufus and P. d. edwardsi had a positive edge response (higher densities in edge habitats), which may be related to edge-related variations in food abundance and quality. Positive edge responses by M. rufus and P. d. edwardsi may ultimately be detrimental due to edge-related anthropogenic factors (e.g., hunting by local people). The negative edge response exhibited by C. major (lower densities in edge habitats) may result from heightened ambient temperatures that inhibit torpor in edge habitats.

Conservation biology of Malagasy strepsirhines: a phylogenetic approach.

The phylogenetic diversity of extant lemurs represents one of the most important but least studied aspects of the conservation biology of primates. The phylogenetic diversity of a species is inversely proportional to the relative number and closeness of its phylogenetic relatives. Phylogenetic diversity can then be used to determine conservation priorities for specific biogeographic regions. Although Malagasy strepsirhines represent the highest phylogenetic diversity among primates at the global level, there are few phylogenetic data on species-specific and regional conservation plans for lemurs in Madagascar. Therefore, in this paper the following questions are addressed for extant lemurs: 1) how does the measure of taxonomic uniqueness used by Mittermeier et al. (1992 Lemurs of Madagascar; Gland, Switzerland: IUCN) equate with an index of phylogenetic diversity, 2) what are the regional conservation priorities based on analyses of phylogenetic diversity in extant lemurs, and 3) what conservation recommendations can be made based on analyses of phylogenetic diversity in lemurs? Taxonomic endemicity standardized weight (TESW) indices of phylogenetic diversity were used to determine the evolutionary component of biodiversity and to prioritize regions for conserving lemur taxa. TESW refers to the standardization of phylogenetic diversity indices for widespread taxa and endemicity of species. The phylogenetic data came from recent genetic studies of Malagasy strepsirhines at the species level. Lemur species were assigned as being either present or absent in six biogeographic regions. TESW indices were combined with data on lemur complementarity and protected areas to assign conservation priorities at the regional level. Although there were no overall differences between taxonomic ranks and phylogenetic rankings, there were significant differences for the top-ranked taxa. The phylogenetic component of lemur diversity is greatest for Daubentonia madagascariensis, Allocebus trichotis, Lepilemur septentrionalis, Indri indri, and Mirza coquereli. Regional conservation priorities are highest for lemurs that range into northeast humid forests and western dry forests. Expansion of existing protected areas in these regions may provide the most rapid method for preserving lemurs. In the long term, new protected areas must be created because there are lemur species that: 1) are not found in existing protected areas, 2) exist only in one or two protected areas, and 3) are still being discovered outside the current network of protected areas. Data on the population dynamics and feeding ecology of phylogenetically important species are needed to ensure that protected areas adequately conserve lemur populations in Madagascar.

Edge effects and their influence on lemur density and distribution in Southeast 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.

Ecogeographic size variations in Sifakas: a test of the resource seasonality and resource quality hypotheses.

Ecogeographic size variations have been documented in some but not all sifakas. Few morphometric or body weight data have been available for two critically endangered subspecies of diademed sifakas: Perrier's sifakas (Propithecus diadema perrieri) and silky sifakas (Propithecus diadema candidus). The objectives of our study were to determine size variations in sifakas and if these variations are related to resource quality and/or resource seasonality. P. d. perrieri and P. d. candidus were captured, weighed, and measured in northern Madagascar. Body weights and morphometrics were compared with other subspecies of diademed sifakas and indris (Indri indri). Differences in body weights and morphometrics between taxa are particularly pronounced for P. d. perrieri compared to P. d. diadema, P. d. edwardsi, and I. indri. Most morphometrics varied in comparisons between P. d. candidus and the other Indriidae (P. d. diadema, P. d. edwardsi, and I. indri). Average body size in sifakas is positively correlated with annual rainfall and negatively correlated with length of dry season. Sifaka body size is not correlated with protein-to-fiber ratios. Thus, size variations in sifakas are related to resource seasonality rather than resource quality. The relationships between the temporal availability of food resources and sifaka body size reflect complex and regionally varying causalities. Detailed, longitudinal information on the ecological factors underlying food selection and nutrient requirements in sifakas are needed to determine the relationship between ecogeographic variables and body size in sifakas.

Dietary patterns in Perrier's sifakas (Propithecus diadema perrieri): a preliminary study.

Some lemur species range into only one habitat type, whereas others range into a variety of habitats. Because plant community structure can differ between habitats, dietary patterns may vary for conspecific groups of primates that range into more than one type of habitat. The goal of our study was to determine how habitat variation influences dietary patterns in Perrier's sifakas (Propithecus diadema perrieri) that range into both dry and riparian forests in northern Madagascar. We collected 542 hr of data on the behavior and diet of two groups of P.d. perrieri from 7 June to 4 August 1998 at Camp Antobiratsy in Analamera Special Reserve, Madagascar. We computed indices of dietary diversity for each group and dietary/plant species similarity between groups. P.d. perrieri in group 1 fed predominantly in dry forest (72.7% of feeding records, n=660), whereas those in group 2 fed most often in riparian forest (73.7% of feeding records, n=666). The index of dietary similarity (0.986) was significantly higher than the index of plant species similarity (0.767). Although the P.d. perrieri in the two study groups fed predominantly in different forest habitats, they ate similar food items in very comparable proportions (but not from the same plant species). However, based on habitat availability measures, neither group fed where they were expected to feed.

Preliminary analysis of drinking from seawater sources by free-ranging rhesus macaques (Macaca mulatta).

Rhesus macaques on Key Lois Island were observed drinking seawater that flowed into four holes they had excavated in the sand. Data were gathered to determine the salinity (TDS) and pH levels of the water and which animals were using the holes. Average TDS level (4,506.8 ± 1,750.8), but not pH (8.1 ± .30) level, of water from the holes differed from the surrounding seawater (TDS = 29,000, pH = 8.0). There were significant variations in TDS and pH levels of water between holes. A total of 249 drinking and 11 digging bouts were observed. Adult females drank and dug most often (46.9% of total bouts). Of the 260 total drinking and digging bouts, 76.1% (N = 198) were concentrated at one hole. This hole had the lowest average TDS level (3,714.2 ± 1,504.4) and one of the highest average pH levels (8.1 ± .29). Age/sex class differences in drinking bout frequencies may have been due to differential social status. We suggest that the holes were excavated to overcome a temporary shortage of provisioned water. © 1993 Wiley-Liss, Inc.