The coexistence of closely related species is not easily understood on the basis of ecological theories. This study investigates the extent of coexistence of two congeneric species of Microcebus murinus (MUR) and M. ravelobensis (RAV) in northwestern Madagascar. Their presence and local relative population densities were determined by capturing and nocturnal transect counts and compared at 22 study sites in the Ankarafantsika National Park. All sites were characterized with regard to their altitude, access to surface water, and 19 structural vegetation characteristics. RAV and MUR were not equally distributed over this regional scale. RAV occurred in more sites and at higher maximum densities than MUR. The relative population densities of both species were significantly and negatively correlated with each other. Whereas the relative population densities of MUR increased with altitude and were highest in dry habitats far from surface water, the relative population densities of RAV generally decreased with altitude and were highest in low altitude habitats close to surface water. The results of the vegetation characteristics also reflect these general trends. The divergent pattern of local and regional coexistence of these two species is discussed and can be best explained either by the existence of a spatially heterogeneous competitive environment or by independent evolutionary pathways in different historic environments. Am. J. Primatol. 71:928–938, 2009. © 2009 Wiley-Liss, Inc.

Varying patterns of coexistence of two mouse lemur species (Microcebus ravelobensis and M. murinus) in a heterogeneous landscape.

Kin-directed food sharing promotes lifetime natal philopatry of both sexes in a population of fish-eating killer whales, Orcinus orca

Philopatry and sex-biased dispersal have a strong influence on population genetic structure, so the study of species dispersal patterns and evolutionary mechanisms shaping them are of great interest. Particularly nongregarious mammalian species present an underexplored field of study: despite their lower levels of sociality compared to group-living species, interactions among individuals do occur, providing opportunities for cryptic kin selection. Among the least gregarious primates are orang-utans (genus: Pongo), in which preferential associations among females have nevertheless been observed, but for which the presence of kin structures was so far unresolved because of the equivocal results of previous genetic studies. To clarify relatedness and dispersal patterns in orang-utans, we examined the largest longitudinal set of individuals with combined genetic, spatial and behavioural data. We found that males had significantly higher mitochondrial DNA (mtDNA) variation and more unique haplotypes, thus underscoring their different maternal ancestries compared to females. Moreover, pedigree reconstruction based on 24 highly polymorphic microsatellite markers and mtDNA haplotypes demonstrated the presence of three matrilineal clusters of generally highly related females with substantially overlapping ranges. In orang-utans and possibly other nongregarious species, comparing average biparental relatedness (r) of males and females to infer sex-biased dispersal is extremely problematic. This is because the opportunistic sampling regime frequently employed in nongregarious species, combined with overlapping space use of distinct matrilineal clusters, leads to a strong downward bias when mtDNA lineage membership is ignored. Thus, in nongregarious species, correct inferences of dispersal can only be achieved by combining several genetic approaches with detailed spatial information.

Parentage-based pedigree reconstruction reveals female matrilineal clusters and male-biased dispersal in nongregarious Asian great apes, the Bornean orang-utans (Pongo pygmaeus).

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.

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

The use of leaf nests has been documented in several mouse lemur species over the last few decades, including the golden-brown mouse lemur. Nest construction, however, has only rarely been observed and detailed descriptions of this process are lacking so far. We aim to determine the relative importance of leaf nests as shelters for the golden-brown mouse lemur, and to test predictions concerning the role of thermoregulation, safety (i.e., protection of infants), and of interspecific competition with the sympatric gray mouse lemurs in regulating nest use. Finally, we intend to clarify whether and how Microcebus ravelobensis constructs the nests, and we provide physical descriptions of seven leaf nests. Nocturnal focal observations were carried out from May 2007 to January 2008 on 18 females, and sleeping sites were regularly monitored during a six-month period. Data were collected from two study sites, one with exclusive presence of M. ravelobensis, and one with co-existence of the two mouse lemur species. Sixty-five out of 379 identified daily sleeping sites were leaf nests. These represented a total of 35 different leaf nests, used by 15 out of 18 females. The relative leaf nest use differed between sites during five out of six months, but without a consistent pattern. Interspecific competition can therefore not explain leaf nest use. Leaf nest use differed seasonally and may be partly explained by thermoregulatory advantages in the site with lower minimum temperatures. Nest use was furthermore higher than expected in both sites during the rearing season that indicates the role of nests in infant protection. For the first time, we could confirm that golden-brown mouse lemurs build leaf nests themselves. Nest building lasted between 46 and 68 min, which shows that this task is time consuming and therefore probably costly. Am. J. Primatol. 72:48–55, 2010. © 2009 Wiley-Liss, Inc.

Leaf nest use and construction in the golden‐brown mouse lemur (Microcebus ravelobensis) in the Ankarafantsika National Park

Summary Natal dispersal reduces the risk of inbreeding, since it is typically biased towards one sex. Golden-brown mouse lemurs {Microcebus ravelobensis) form mixed-sex sleeping groups whose genetic composition is not yet known. This study investigates the composition of eight sleeping groups and the dispersal pattern in a free-living population of this nocturnal primate. Genetic relatedness and parentage was calculated for 101 individuals (43 males, 58 females) based on microsatellite analyses. Matrilinear relatedness played a major role in the composition of sleeping groups. Co-sleepers were significantly more closely related than members of different sleeping groups. Dispersal was not strongly biased towards one sex. More than 50% of all potential sons (N = 19) and daughters (N = 26) remained in their natal range well into their first mating season, but potential sons disappeared more frequently than daughters lateron. These data indicate that many sons may have delayed their dispersal. This delayed dispersal may have increased the risk of inbreeding, since closely related males and females stayed within one home range diameter of each other quite regularly in the mating season. In fact, two definite cases of inbreeding were detected. The likely benefits of delayed dispersal are discussed in view of the confirmed cases of inbreeding.

E. Zimmermann

Papers

Sociogenetic structures, dispersal and the risk of inbreeding in a small nocturnal lemur, the golden-brown mouse lemur (Microcebus ravelobensis)