The last decade's lemur research includes successes in discovering new living and extinct species and learning about the distribution, biogeography, physiology, behavior, and ecology of previously little-studied species. In addition, in both the dry forest and rain forest, long-term studies of lemur demography, life history, and reproduction, have been completed in conjunction with data on tree productivity, phenology, and climate. Lemurs contrast with anthropoids in several behavioral features, including female dominance, targeted female-female aggression, lack of sexual dimorphism regardless of mating system, sperm competition coupled with male-male aggression, high infant mortality, cathemerality, and strict seasonal breeding. Hypotheses to explain these traits include the "energy conservation hypothesis" (ECH) suggesting that harsh and unpredictable climate factors on the island of Madagascar have affected the evolution of female dominance, and the "evolutionary disequilibrium hypotheses" (EVDH) suggesting that the recent megafauna extinctions have influenced lemurs to become diurnal. These hypotheses are compared and contrasted in light of recent empirical data on climate, subfossils, and lemur behavior. New data on life histories of the rain forest lemurs at Ranomafana National Park give further support to the ECH. Birth seasons are synchronized within each species, but there is a 6-month distribution of births among species. Gestation and lactation lengths vary among sympatric lemurs, but all lemur species in the rain forest wean in synchrony at the season most likely to have abundant resources. Across-species weaning synchrony seen in Ranomafana corroborates data from the dry forest that late lactation and weaning is the life history event that is the primary focus of the annual schedule. Lemur adaptations may assure maximum offspring survival in this environment with an unpredictable food supply and heavy predation. In conclusion, a more comprehensive energy frugality hypothesis (EFH) is proposed, which postulates that the majority of lemur traits are either adaptations to conserve energy (e.g., low basal metabolic rate (BMR), torpor, sperm competition, small group size, seasonal breeding) or to maximize use of scarce resources (e.g., cathemerality, territoriality, female dominance, fibrous diet, weaning synchrony). Among primates, the isolated adaptive radiation of lemurs on Madagascar may have been uniquely characterized by selection toward efficiency to cope with the harsh and unpredictable island environment.

Lemur traits and Madagascar ecology: coping with an island environment.

Via the current model on the evolutionary ecology of female social relationships, Sterck et al. (1997) argue that ecological conditions determine how competition over food resources affects female fitness. The relative importance of different modes of competition then affects female social relationships and dispersal patterns. I outline the model and review relevant data. There are 3 modes of feeding competition: within-group scramble (WGS), within-group contest (WGC), and between-group contest (BGC), which occur in various combinations in different populations of nonhuman primates. Ecological measures support predictions that limiting resources lead to WGS and clumped resources induce WGC. The ecological basis of BGC remains elusive, but it is probably linked to resource abundance. Tests of the proxies of feeding competition support the idea that short-term search substrates and increasing group size lead to WGS, while high-quality patches of intermediate size relative to group size lead to WGC. However, when tested across populations, independent measures of aggression rates do not always match the actual or presumed competitive regimes. This mismatch might be explained by confounding factors and the predominately indirect measures of feeding competition. Predicted relationships between feeding competition and female social relationships/dispersal are only partly supported. This might be attributed to the fact that few studies have taken ultimate approaches using mechanistic correlates of fitness (net energy gain) or lifetime reproductive success to measure consequences of feeding competition. But to resolve existing inconsistencies, additional factors need to be taken into account as well, for example, male sexual strategies may affect female feeding competition; constraints on group size may enforce female dispersal; and demography may alter rates of alliances. More explicitly, ultimate approaches are needed to test the consistency of the socioecological model.

Competition for resources and its behavioral consequences among female primates

The lemurs of Madagascar are a unique radiation of primates that show an extraordinary diversity of lifestyles, morphologies and behaviours. However, very little is known about the relative antiquity of lemuriform clades due to the lack of terrestrial fossils for the Tertiary of Madagascar. Here, we employ a Bayesian method to estimate divergence dates within the lemuriform radiation using several unlinked gene loci and multiple fossil calibrations outside the lemuriform clade. Two mitochondrial genes (cytochrome oxidase II and cytochrome b ), two nuclear introns (transthyretin intron 1 and von Willebrand factor gene intron 11) and one nuclear exon (interphotoreceptor retinoid binding protein, exon 1) are used in separate and combined analyses. The genes differ in taxon sampling and evolutionary characteristics but produce congruent date estimates. Credibility intervals narrow considerably in combined analyses relative to separate analyses due to the increased amount of data. We also test the relative effects of multiple vs. single calibration points, finding that, when only single calibration points are employed, divergence dates are systematically underestimated. For the mitochondrial DNA data set, we investigate the effects of sampling density within the mouse lemur radiation (genus Microcebus ). When only two representative species are included, estimated dates throughout the phylogeny are more recent than with the complete-species sample, with basal nodes less affected than recent nodes. The difference appears to be due to the manner in which priors on node ages are constructed in the two analyses. In nearly all analyses, the age of the lemuriform clade is estimated to be approximately 62–65 Ma, with initial radiation of mouse lemurs and true lemurs (genus Eulemur ) occurring approximately 8–12 Ma. The antiquity of the mouse lemur radiation is surprising given the near uniform morphology among species. Moreover, the observation that mouse lemurs and true lemurs are of similar ages suggests discrepancies in rates of morphological, behavioural and physiological evolution in the two clades, particularly with regard to characteristics of sexual signalling. These differences appear to correlate with the nocturnal vs. diurnal lifestyles, respectively, of these two primate groups.

/pdf/divergence-dates-for-malagasy-lemurs-estimated-from-multiple-49kra9xata.pdf

Divergence dates for Malagasy lemurs estimated from multiple gene loci: geological and evolutionary context

Phylogenetic analysis of mtDNA sequence data confirms the observation that species diversity in the world's smallest living primate (genus Microcebus) has been greatly underestimated. The description of three species new to science, and the resurrection of two others from synonymy, has been justified on morphological grounds and is supported by evidence of reproductive isolation in sympatry. This taxonomic revision doubles the number of recognized mouse lemur species. The molecular data and phylogenetic analyses presented here verify the revision and add a historical framework for understanding mouse lemur species diversity. Phylogenetic analysis revises established hypotheses of ecogeographic constraint for the maintenance of species boundaries in these endemic Malagasy primates. Mouse lemur clades also show conspicuous patterns of regional endemism, thereby emphasizing the threat of local deforestation to Madagascar's unique biodiversity.

https://www.pnas.org/content/pnas/97/21/11325.full.pdf

Remarkable species diversity in Malagasy mouse lemurs (primates, Microcebus)

The genus Microcebus (mouse lemurs) are the smallest extant primates. Until recently, they were considered to comprise two different species: Microcebus murinus, confined largely to dry forests on the western portion of Madagascar, and M. rufus, occurring in humid forest formations of eastern Madagascar. Specimens and recent field observations document rufous individuals in the west. However, the current taxonomy is entangled due to a lack of comparative material to quantify intrapopulation and intraspecific morphological variation. On the basis of recently collected specimens of Microcebus from 12 localities in portions of western Madagascar, from Ankarana in the north to Beza Mahafaly in the south, we present a revision using external, cranial, and dental characters. We recognize seven species of Microcebus from western Madagascar. We name and describe 3 spp., resurrect a previously synonymized species, and amend diagnoses for Microcebus murinus (J. F. Miller, 1777), M. myoxinus Peters, 1852, and M. ravelobensis Zimmermann et al., 1998.

/pdf/taxonomic-revision-of-mouse-lemurs-microcebus-in-the-western-1uussh2uoi.pdf

Taxonomic Revision of Mouse Lemurs (Microcebus) in the Western Portions of Madagascar

Sleeping sites are a potentially important resource for grey mouse lemurs since they are confronted with high daily temperature fluctuations and a high predation pressure. In order to determine the existence and degree of resource competition, sleeping site characteristics, locations, and usage patterns as well as sleeping group compositions were investigated in a 3 month field study in a dry deciduous forest of northwestern Madagascar. The daily sleeping sites of females were on average better insulated and safer than those of males. Males used more sleeping sites and changed the site more often than females. During the whole study, males slept alone, whereas the females formed stable sleeping groups in on average 83.7% of the days. Sex-specific differences in usage patterns might be explained by intersexual resource competition and female dominance and could possibly be related to differential parental investment of the sexes. The underlying study indicates that sleeping sites may be a restricted and defendable resource for grey mouse lemurs. The investigation gives new insights into the distribution patterns and social organization of this species.

Sex-specific usage patterns of sleeping sites in grey mouse lemurs (Microcebus murinus) in northwestern Madagascar.

0015–5713/98/0692–0106$15.00/0 E-Mail karger@karger.ch Fax+ 41 61 306 12 34 This article is accessible online at: http://www.karger.ch http://BioMedNet.com/karger Prof. Elke Zimmermann Institute of Zoology, Tierarztliche Hochschule Hannover Bunteweg 17, D–30559 Hannover (Germany) Tel. 0511 953 8740, Fax 0511 953 8586 E-Mail ezimmer@zoologie.tiho-hannover.de Received: December 5, 1996 Accepted: January 3, 1997 Folia Primatol 1998;69:106–114 Brief Report

/pdf/sympatric-mouse-lemurs-in-north-west-madagascar-a-new-rufous-2lbzltavrt.pdf

Sympatric mouse lemurs in north-west Madagascar: a new rufous mouse lemur species (Microcebus ravelobensis).

https://epub.ub.uni-muenchen.de/16729/1/10_1159_000021571.pdf

Sabine Cepok

Papers

Sex-specific usage patterns of sleeping sites in grey mouse lemurs (Microcebus murinus) in northwestern Madagascar.

Sympatric mouse lemurs in north-west Madagascar: a new rufous mouse lemur species (Microcebus ravelobensis).

oooooooooooooooooooooooooooooooo Sympatric Mouse Lemurs in North-West Madagascar: A New Rufous Mouse Lemur Species (Microcebus ravelobensis)