Habitat Characteristics and Lemur Species Richness in Madagascar1

In the extant lemur communities of Madagascar the number of lemur species increases with increasing number of tree species. In forests with few tree species lemurs use patches with higher number of tree species than average. However, in forest plots with high number of tree species, lemurs stay in places with lower number of tree species than average. At low tree species diversity a minimum number of different tree species seems to be required within the animals' home range to assure year-round food availability. At high tree species diversity tree species essential for survival might be diluted by resources which are of no use for lemurs, thus increasing energetic expenses for traveling between suitable patches. According to the present analyses, structural diversity is of subordinate importance to the number of tree species as a correlate of lemur species richness. Within limits of disturbance intensity and on a small geographic scale, disturbances increase forest productivity. Lemurs reach higher species numbers and population densities in slightly disturbed areas compared with undisturbed sites. This peaked curve of the number of lemur species over disturbance, however, may not only be a consequence of “resource dilution” in undisturbed sites and higher food abundance in slightly disturbed areas, but also a consequence of selective extinction of lemur species which were unable to cope with the disturbance regime exaggerated by human interference over the last few hundred or thousand years.     

Community monopolization: local adaptation enhances priority effects in an evolving metacommunity

The diversity and composition of biological communities might often depend on colonization history because early colonists can exclude future colonists through a priority effect. These priority effects, which have been observed across a wide variety of ecosystems, often arise because early colonists have sufficient time to use available resources efficiently and subsequently withhold them from invaders. Here, we explore the extent to which rapid local adaptive evolution contributes to the pervasiveness of these priority effects. Using an individual-based simulation, we show that early colonization allows the descendants of colonists to adapt to novel conditions and reduce the establishment success of an initially ecologically equivalent competing species. Our model predicts that slight differences in colonization timing and adaptive capacity between species can substantially alter the dynamics and diversity of communities. We also show that priority effects and gene flow can generate a novel mechanism for the expansion and retraction of species distributions in a metacommunity. Our results suggest that local adaptation combined with stochastic colonization events can obscure direct relationships between species distributions and environmental gradients. Given the increasing recognition of rapid, microgeographic evolution in natural populations, we expect that evolutionary priority effects could affect the structure and dynamics of many natural metacommunities.     

Frugivory and Seed Dispersal by Four Species of Primates in Madagascar's Eastern Rain Forest1

In the unique faunal assemblage of the Malagasy rain forest, lemurs appear to play particularly important roles as seed-dispersing frugivores. A three-month study of feeding ecology and seed dispersal by four species of lemurs in Madagascar's eastern rain forest found that three species, Eulemur rubriventer, Eulemur fulvus, and Varecia variegata were seed dispersers, and the fourth, Propithecus diadema, was a seed predator. In germination trials, seeds passed by lemurs sprouted significantly faster and in greater numbers than those not passed by lemurs. Analysis of fruit morphologies of 69 local plant taxa producing fleshy fruits during the study period found that these fruits fell into two well-defined color categories that correlated significantly with fruit size. Seventy seven percent of fleshy fruits greater than 10 mm in diameter were colored green, brown, tan, purplish, or black, while all fruits less than 10 mm in diameter were colored red, yellow, orange, pink, blue, or white. Three introduced exotic plant species provided exceptions to this pattern, producing fruits which were larger than 10 mm and pink or orange. Fruits chosen by the primates in this study were usually larger than 10 mm in diameter and were in nearly all cases colored green, brown, tan, purplish, red, or some combination of these colors. Morphological traits shared by fruits of multiple plant taxa in the diets of seed-dispersing lemurs suggest possible coevolved relationships between Malagasy rain forest plants and lemurs.     

Lemur Diversity in Madagascar

A basic understanding of the taxonomy, diversity, and distributions of primates is essential for their conservation. This review of the status of the taxonomy of lemurs is based on a 5-d workshop entitled “Primate Taxonomy for the New Millennium,” held at the Disney Institute, Orlando, Florida, in February 2000. The aim is not to present a taxonomic revision, but to review our current understanding of the diversity and current and past ranges of lemurs and indicate where there is controversy, discrepancy, or lack of knowledge. Our goal therefore is to provide a baseline for future taxonomic investigation, as well as a clearer focus for research and conservation priorities. We here focus on the lemurs of Madagascar and recognize 5 families, 15 genera, and 99 species and subspecies. We list 39 species of lemurs described since 2000: 2 dwarf lemurs, Cheirogaleus; 11 mouse lemurs, Microcebus; a giant mouse lemur, Mirza; a bamboo lemur, Hapalemur; 17 sportive lemurs, Lepilemur; and 7 woolly lemurs, Avahi. Taxonomic revisions have resulted in the resurrection of a further 9 taxa. However, the figures do not represent the total diversity of Malagasy lemurs because more new species are being identified via new field studies and accompanying genetic research, and should be described in the near future.     

Direct and Indirect Impacts of Raptor Predation on Lemurs in Southeastern Madagascar

I calculated rates of predation by 2 species of diurnal raptors, Polyboroides radiatus and Accipiter henstii, on the lemur community of Ranomafana National Park, Madagascar from 2700 h of observation and 470 prey deliveries at 7 nests of each hawk species. The 2 hawks consumed 7 of 12 lemurs found in the park region, with a body mass of 63–3500 g and including diurnal and nocturnal species of all group sizes. Calculations of predation rates indicate that raptor predation is a significant cause of mortality for lemur populations relative to other causes. Minimum rates of predation by Polyboroides radiatus and Accipiter henstii on Microcebus rufus, Cheirogaleus major, Avahi laniger, Hapalemur griseus, Eulemur fulvus rufus, Eulemur rubriventer, and Varecia variegata resulted in the raptors removing of 1–21% of the population per yr, similar to other rates of predation on primates documented in the literature. Modeling of lemur populations under varying levels of raptor predation pressure that I calculated found that one may attribute 3–17% of adult, juvenile, and infant mortality for nocturnal lemurs and 2–66% of adult, juvenile, and infant mortality for diurnal lemurs to diurnal raptor predation. Raptor predation may significantly depress intrinsic growth rates and carrying capacity of Avahi laniger, Hapalemur griseus, Eulemur fulvus rufus, Eulemur rubriventer, and Varecia variegata owing to their low fecundities, long life spans, and long age to sexual maturation. Nocturnal lemurs may best avoid predation by diurnal raptors by exhibiting a solitary lifestyle and cryptic antipredator tactics, whereas, diurnal lemurs benefit less by increasing group size than by using specific antipredator tactics.     

Dental use wear in extinct lemurs: evidence of diet and niche differentiation

A new technique for molar use-wear analysis is applied to samples of all 16 species of extinct lemurs with known dentitions, as well as to a large comparative sample of extant primates. This technique, which relies on the light refractive properties of wear pits and scratches as seen under a standard stereoscopic microscope, has shown itself to be effective in distinguishing the diets of ungulates and extant primates. We draw dietary inferences for each of the 16 extinct lemur species in our database. There is a strong phylogenetic signal, with the Palaeopropithecidae showing use-wear signatures similar to those of the Indriidae; extinct lemurids (Pachylemur spp.) showing striking similarities to extant lemurids (except Hapalemur spp.); and Megaladapis showing similarities to Lepilemur spp. Only the Archaeolemuridae have dietary signatures unlike those of any extant lemurs, with the partial exception of Daubentonia. We conclude that the Archaeolemuridae were hard-object feeders; the Palaeopropithecidae were seed predators, consuming a mixed diet of foliage and fruit to varying degrees; Pachylemur was a fruit-dominated mixed feeder, but not a seed predator; and all Megaladapis were leaf browsers. There is no molar use wear evidence that any of the extinct lemurs relied on terrestrial foods (C4 grasses, tubers, rhizomes). This has possible implications for the role of the disappearance of wooded habitats in the extinction of lemurs.     

Madagascar's Lemurs: Cryptic diversity or taxonomic inflation?

We live in inflationary times. A quarter of a century ago, cigarettes were about $1 a pack in New York City, a bottle of Château Beaucastel set you back $15, and there were 36 different species of lemur alive in Madagascar 1 (Table 1). Today the equivalent figures are $7.40, $95, and 83 lemur species 2 (Table 1). The increase in dollar prices has a lot to do with the economics of growth, something that obviously cannot be sustained indefinitely on a finite planet. Is the recent inflation in lemur taxonomy any more secure? The question is all the more worth asking because this is no isolated phenomenon: Madagascar's primates have not been alone in multiplying. The same burgeoning of species names has occurred throughout the order Primates 3 , 4 and beyond, 5 provoking both concern and energetic debate. 6 - 9 Interestingly, this debate has largely unfolded among ecologists, conservationists, and other “consumers” of taxonomy; many “producers” seem to be content to generate new taxonomies with a remarkable lack of introspection, as if they were no more than passive consequences of more lofty concerns. And because the same causes underlie taxonomic inflation in Madagascar as elsewhere, this extraordinary island once again presents us with a microcosm of the larger world.     

Resting metabolic rates of Lepilemur ruficaudatus

Seventeen sportive lemurs (Lepilemur ruficaudatus) were captured in a dry, deciduous forest in western Madagascar, and the resting metabolic rates were measured. According to the data, resting metabolic rates of Lepilemur ruficaudatus are among the lowest of mammalian folivores recorded so far. © 1996 Wiley-Liss, Inc.     

Elevational Ranges of Lemurs in the Humid Forests of Madagascar

The geological history of Madagascar contains events that may have given rise to unusual factors that molded the primate communities of the island. In order to understand the impact of the factors we reviewed the elevational distribution of extant primate species in the humid forests. The elevational structuring of primates in the humid forests of Madagascar shows considerable differences from other tropical areas in the New World and Asia, which is correlated with the lack of large tracts of humid lowland habitat on Madagascar since it split from India.