Factors influencing terrestriality in primates of the Americas and Madagascar.

Among mammals, the order Primates is exceptional in having a high taxonomic richness in which the taxa are arboreal, semiterrestrial, or terrestrial. Although habitual terrestriality is pervasive among the apes and African and Asian monkeys (catarrhines), it is largely absent among monkeys of the Americas (platyrrhines), as well as galagos, lemurs, and lorises (strepsirrhines), which are mostly arboreal. Numerous ecological drivers and species-specific factors are suggested to set the conditions for an evolutionary shift from arboreality to terrestriality, and current environmental conditions may provide analogous scenarios to those transitional periods. Therefore, we investigated predominantly arboreal, diurnal primate genera from the Americas and Madagascar that lack fully terrestrial taxa, to determine whether ecological drivers (habitat canopy cover, predation risk, maximum temperature, precipitation, primate species richness, human population density, and distance to roads) or species-specific traits (body mass, group size, and degree of frugivory) associate with increased terrestriality. We collated 150,961 observation hours across 2,227 months from 47 species at 20 sites in Madagascar and 48 sites in the Americas. Multiple factors were associated with ground use in these otherwise arboreal species, including increased temperature, a decrease in canopy cover, a dietary shift away from frugivory, and larger group size. These factors mostly explain intraspecific differences in terrestriality. As humanity modifies habitats and causes climate change, our results suggest that species already inhabiting hot, sparsely canopied sites, and exhibiting more generalized diets, are more likely to shift toward greater ground use.

Frugivory in four sympatric lemurs: implications for the future of Madagascar's forests.

Although some conservationists accept that not all species can be saved, we illustrate the difficulty in deciding which species are dispensable. In this article, we examine the possibility that the integrity of a forest relies on its entire faunal assemblage. In Madagascar, one faunal group, the lemurs, accounts for the greatest biomass and species richness among frugivores. For example, 7 of the 13 sympatric lemur species in Madagascar's eastern rainforests consume primarily fruit. Because of this, we suggest that some tree species may rely heavily on particular lemur taxa for both seed dispersal and germination. In Ranomafana National Park, the diets for four of the day-active lemur frugivores have been documented during annual cycles over a 5-year period. We predicted that, although the fruit of some plant taxa would be exploited by multiple lemur species, the fruit of others would be eaten by one lemur species alone. Analyses reveal that while lemurs overlap in a number of fruit taxa exploited, 46% (16/35) of families and 56% (29/52) of genera are eaten exclusively by one lemur species. We, therefore, predict local changes in forest composition and structure if certain of these lemur species are eliminated from a forest owing to hunting, disease, or habitat disturbance. We also suggest that this result may be of global significance because carbon sequestration by the tropical forests in Madagascar may be reduced as a result of this predicted change in forest composition.

Population demography and social structure changes in Eulemur fulvus rufus from 1988 to 2003.

Eulemur fulvus rufus has been described as having stable multi-male/multi-female groups, a male-biased sex ratio, and female philopatry. However, in a 16-year study of this subspecies we documented a great deal of demographic change as several groups permanently fissioned, some groups disappeared, and new groups formed. We split the dataset into two periods, 1988 to 1993 and 1994 to 2003, which coincided with the first disappearance of a study group (in August 1994) and the first permanent group fission (in December 1994). The average group size decreased by nearly half between the study periods (10.5-5.6), while the frequency of group membership changes increased (2.0-8.3 times/year), and the birth rate decreased (0.56-0.38). Females, as well as males, immigrated into study groups and transferred between groups, something that has been rarely seen in this subspecies. We also found a significant decline in the amount of fruit from the earliest part of the study to the latter part of the study. Study groups did not switch to other types of foods during periods of fruit shortage, but traveled outside of their home range areas more often over the study period. Finally, the density E. f. rufus decreased in the study area while the densities of their main food competitors, Varecia variegata and Eulemur rubriventer, increased. Although few primate populations are numerically stable over time, we suggest that female behavioral responses to decreases in fruit availability may have influenced some of the demographic changes we witnessed in this study.

Spatial memory during foraging in prosimian primates: Propithecus edwardsi and Eulemur fulvus rufus.

A variety of anthropoids travel efficiently from one food source to another, although there is disagreement over how this is accomplished over large-scale space. Mental maps, for example, require that animals internally represent space, geometrically locate landmarks, use true distance and direction, and generate novel shortcuts to resources. Alternately, topological or route-based maps are based on a network of fixed points, landmarks and routes so that one food patch can be linked with another. In this study we describe travel patterns between food sources for two prosimian species found in southeastern Madagascar, Propithecus edwardsi and Eulemur fulvus rufus. Both species are dependent on fruit and have large home range sizes. By comparing interpatch distances, patch size and turning angles, we found that both species prefer nearest neighbor food patches and P. edwardsi travels in relatively straight lines. The amount of backtracking seen in E. f. rufus may be linked to their large group size and dependence on large-crowned fruit trees. We suggest that the goal-oriented foraging of both prosimian species is dependent on a topological or route-based map. These are rare behavioral data relevant to ecological and social contexts of primate cognitive evolution.

Behavioral development of captive male hybrid cercopithecine monkeys.

In this study, we compare the behavioral development of captive male vervet monkeys (Chlorocebus pygerythrus) and Sykes' monkeys (Cercopithecus albogularis) to male hybrids of these species. Focal animal sampling sessions were conducted on the study subjects from birth to 90 months of age. Behavioral categories (affiliative, approach, play, sexual, tension, aggressive) were plotted for each species and the hybrids; curves were fitted using polynomial regression and were evaluated with the Wilcoxon signed ranks test. With the exception of play behaviors, the male C. pygerythrus x C. albogularis hybrids modeled the Sykes' pattern of behavioral development. We suggest that this result reflects a sex-linked paternal effect.

Does female dominance facilitate feeding priority in black-and-white ruffed lemurs (Varecia variegata) in southeastern Madagascar?

Although many Malagasy lemurs are thought to be female dominant and to have female feeding priority, to date the relationship between these behaviors has been rigorously established only in Lemur catta, and other ways that females might achieve feeding priority have not been examined closely. Erhart and Overdorff [International Journal of Primatology 20:927-940, 1999] suggested that one way female primates achieve feeding priority is to initiate and lead groups to food, thereby gaining access to the food first and positively influencing their food intake compared to other group members. Here we describe female dominance patterns and potential measures of feeding priority in two groups of black-and-white ruffed lemurs (Varecia variegata) that were observed over a 15-month period in southeastern Madagascar. We predicted that the females would 1) be consistently dominant to males, 2) lead groups to food sources more often than males, and 3) have higher feeding rates compared to males when they arrived at food sources first. The results were dissimilar between the study groups. During the study, the oldest adult female in group 1 died. There was no evidence for female dominance in this group, and the remaining (likely natal) female did not lead the group more often, nor did she have a higher food intake than males. Group 1 dispersed shortly after the time frame reported here. In contrast, the resident female in group 2 was dominant to group males (based on agonistic interactions), led the group to food sources more often, and experienced a higher food intake when she arrived first at a food source. How these patterns vary over time and are influenced by the number of females in groups, group stability, food quality, and reproductive condition will be examined in future analyses.