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.

Dietary preferences and feeding strategies of Colombian highland woolly monkeys.

Primates are very selective in the foods they include in their diets with foraging strategies that respond to spatial and temporal changes in resource availability, distribution and quality. Colombian woolly monkeys (Lagothrix lagotricha lugens), one of the largest primate species in the Americas, feed mainly on fruits, but they also eat a high percentage of arthropods. This differs from closely related Atelid species that supplement their diet with leaves. In an 11 month study, we investigated the foraging strategies of this endemic monkey and assessed how resource availability affects dietary selection. Using behavioural, phenological, arthropod sampling and metabarcoding methods, we recorded respectively foraging time, forest productivity, arthropod availability in the forest and arthropod consumption. Scat samples and capturing canopy substrates (i.e. moss, bromeliads, aerial insects) were used for assigning arthropod taxonomy. The most important resource in the diet was fruits (54%), followed by arthropods (28%). Resource availability predicted feeding time for arthropods but not for fruits. Further, there was a positive relationship between feeding time on fruits and arthropods, suggesting that eating both resources during the same periods might work as an optimal strategy to maximize nutrient intake. Woolly monkeys preferred and avoided some fruit and arthropod items available in their home range, choosing a wide variety of arthropods. Geometrid moths (Lepidoptera) were the most important and consistent insects eaten over time. We found no differences in the type of arthropods adults and juveniles ate, but adults invested more time foraging for this resource, especially in moss. Although woolly monkeys are generalist foragers, they do not select their food items randomly or opportunistically.

Influence of Arthropod and Fruit Abundance on the Dietary Composition of Highland Colombian Woolly Monkeys (Lagothrix lagotricha lugens).

The woolly monkey's (Lagothrix lagotricha) diet consists primarily of fruits, and for some populations fruit availability predicts its frequency in the diet. Woolly monkeys also eat new leaves, flowers, seeds and arthropods, but it is unclear whether arthropods are consumed in proportion to their environmental availability. We tested the hypothesis that arthropod consumption by woolly monkeys depends on availability. We studied a group of woolly monkeys for 10 months in 2013-2014, in Cueva de los Guácharos National Park, Colombia, in order to test the hypothesis that arthropod consumption by woolly monkeys depends on availability. We carried out surveys of plant phenology each month for 10 months to estimate fruit productivity, and every 2 months we also surveyed 3 canopy substrates (leaves, mosses and bromeliads) to quantify variation in arthropod biomass. During this time, we also quantified the diet of a woolly monkey group. The items most consumed were fruits (60%), followed by arthropods (24%), leaves (13%) and miscellaneous other items (3%). Arthropod biomass in the canopy did not vary considerably over the 10 months (0.014-0.037 g/g substrate) but was lower at the end of the second rainy season. A positive correlation was found between availability and entomophagy, but only when arthropods were relatively abundant. We did not find a relationship between arthropod and fruit feeding frequencies. Our results indicate that arthropods are prevalent in the diet of the woolly monkeys due to their abundance in Andean forest canopies.