On the scaling of activity in tropical forest mammals

Activity range – the amount of time spent active per day – is a fundamental aspect contributing to the optimization process by which animals achieve energetic balance. Based on their size and the nature of their diet, theoretical expectations are that larger carnivores need more time active to fulfil their energetic needs than do smaller ones and also more time active than similar‐sized non‐carnivores. Despite the relationship between daily activity, individual range and energy acquisition, large‐scale relationships between activity range and body mass among wild mammals have never been properly addressed. This study aimed to understand the scaling of activity range with body mass, while controlling for phylogeny and diet. We built simple empirical predictions for the scaling of activity range with body mass for mammals of different trophic guilds and used a phylogenetically controlled mixed model to test these predictions using activity records of 249 mammal populations (128 species) in 19 tropical forests (in 15 countries) obtained using camera traps. Our scaling model predicted a steeper scaling of activity range in carnivores (0.21) with higher levels of activity (higher intercept), and near‐zero scaling in herbivores (0.04). Empirical data showed that activity ranges scaled positively with body mass for carnivores (0.061), which also had higher intercept value, but not for herbivores, omnivores and insectivores, in general, corresponding with the predictions. Despite the many factors that shape animal activity at local scales, we found a general pattern showing that large carnivores need more time active in a day to meet their energetic demands. Introduction Activity range – the amount of time, in hours, spent active per day – is a fundamental outcome of the complex physiological and behavioral optimization process by which animals ensure that energy input keeps pace with energy output. In addition to basal metabolism, animals face costs of foraging, acquiring mates and shelter, building reserves for lean times and escaping predators (Carbone et al. 2007, Halle and Stenseth 2012). Environmental and ecological factors that vary through the day (e.g. luminosity, temperature, predation risk and competition avoidance) constrain activity to certain times, depending on morpho‐physiological limitations (Castillo‐Ruiz et al. 2012, Hut et al. 2012). In addition, animals need time to rest in order to recover their cognitive or physical condition (Siegel 2005). Thus, they must optimize their activity range to meet their resource requirements, while dealing with natural daily cycles and saving time for sleep/rest (Downes 2001, Siegel 2005, Cozzi et al. 2012). The resource requirements of mammals are related to basal metabolic rate, which scales positively with body mass (Kleiber 1932, Isaac and Carbone 2010), while predation risk decreases with body mass (Sinclair et al. 2003, Hopcraft et al. 2009). Because high predation risk constrains activity while high resource needs increases activity range (Cozzi et al. 2012, Suselbeek et al. 2014), the question arises whether and how activity range also scales with body mass. Day range (total distance travelled in a day) and home range (area in which animals perform their daily activities) scales positively with body mass and are key metrics to understand the resource requirements of an animal (McNab 1963, Kelt and Van Vuren 2001, Carbone et al. 2005, Tamburello et al. 2015). As activity range is related to space‐use metrics (i.e. home range and day range), it is hence, also related to the acquisition of energy. Given that, one might expect activity range to increase with body mass. However, we have a poor understanding of how this relationship actually looks. Previous work developed predictions of body mass scaling with day range (Garland 1983, Carbone et al. 2005) and travel speed (Carbone et al. 2007, Rowcliffe et al. 2016). From a simple physical viewpoint, activity range should equal the day range divided by average travel speed. It should thus be possible to infer the scaling of activity range with body mass from these relationships. Some of the variation in space use across species that is not explained by body mass is associated with different evolutionary histories and ecological traits (McNab 1963, Kelt and Van Vuren 2001, Price and Hopkins 2015, Tamburello et al. 2015). Diet is the most conspicuous of these, because primary and secondary productivity present different overall yields and accessibility for consumers (Jetz et al. 2004), which in turn influence individual movements (Carbone et al. 2005) and potentially activity range, when exploiting resources at different trophic levels. The nature of the diet aggravates the higher energetic demands of larger carnivores. Predators have considerable energetic constraints related to hunting and handling their prey (Gorman et al. 1998, Carbone et al. 1999) as animal prey can be rare, widely dispersed, unpredictable in time and space and not storable (Jetz et al. 2004, Carbone et al. 2007). Therefore, carnivores have the lowest energy supply rates (supply rate of usable resources available inside the home range), independent of body mass, when compared to other diet categories (Jetz et al. 2004) besides exploring larger areas and traveling greater daily distances (McNab 1963, Kelt and Van Vuren 2001, Carbone et al. 2005, Tamburello et al. 2015). Therefore, larger animals occupy larger areas than small ones, and carnivores occupy larger areas than do similar‐sized non‐carnivores (Jetz et al. 2004, Tamburello et al. 2015). To date, few studies have considered interspecific variation in activity range with body mass and other species traits. For example, van Schaik and Griffiths (1996) and Gómez et al. (2005) anecdotally suggested that larger mammal species are cathemeral (i.e. active day and night), which implies that they can be active during a larger proportion of the 24‐h cycle. Rowcliffe et al. (2014) found that activity range is positively correlated with body mass in tropical forest mammals in Panama. Ramesh et al. (2015) found a negative relationship between body mass and activity concentration (i.e. how concentrated in few hours is the activity of an animal during the day) in Indian mammals, also equating to a positive association between activity range and body mass. However, no study has explored variation in activity range across a diverse range of species, while controlling for phylogeny and diet. This has been, at least in part, due to a lack of consistent data available on a wide range of species. Recent work using camera traps (Oliveira‐Santos et al. 2013, Rowcliffe et al. 2014), however, has demonstrated that accurate estimates of activity range can be obtained from photographic records from camera traps. Given the large and rapidly increasing volume of camera‐trapping data available globally (Burton et al. 2015), these approaches, consistently applied across a wide range of studies, can provide an important basis for the large‐scale study of activity. Here, we provided simple empirical predictions for the scaling of activity range with body mass for mammals of different trophic guilds. To test these predictions, we estimated the activity range for 249 populations of 128 terrestrial mammal species across 19 tropical forests, and used a phylogenetically controlled mixed model to determine how activity range scales with body mass by diet. As larger animals occupy larger areas than small ones, and carnivores occupy larger areas than do similar‐sized non‐carnivores (Jetz et al. 2004), we hypothesize that carnivores will present a higher scaling of activity range with body mass and also higher activity ranges for a given mass (higher intercept) when compared to herbivores, omnivores and insectivores

Parapatric pied and red-handed tamarin responses to congeneric and conspecific calls

Aggressive behaviors are widespread among territorial species and asymmetrical aggressiveness may imply differential access to resources. At a larger scale, such asymmetry may also drive shifts in geographic distributions. The pied tamarin (Saguinus bicolor) is an endangered Amazonian primate species with a small natural range. In recent decades further reduction of its range has been observed coincident with the expansion of the range of the red-handed tamarin's (Saguinus midas), which appears to be encroaching into the area otherwise occupied by the pied tamarin. Here we test if, at range boundaries, red-handed tamarin produces more aggressive vocalizations than the pied tamarin. We performed a series of 96 playback trials presenting both congeneric and conspecific long calls to sixteen groups of red-handed tamarins and fourteen of pied tamarins. We recorded their territorial, agonistic, alarm vocalizations, and the duration of their calling displays after broadcasts. In doing so, we assessed whether agonistic displays were more likely to occur in response to congeneric than conspecific calls in areas of sympatry. We found that the red-handed tamarin was more likely to emit more territorial calls when listening to pied tamarins than to its own species in sympatric areas, but found no differences in vocal responses from either species in relation to agonistic calls or duration of display in sympatric and allopatric areas. Furthermore, the red-handed tamarin emitted more alarm calls when listening to pied tamarin, independently of the geographic circumstances. Overall, we found that acoustic displays may be mediating species interaction in areas of sympatry. Together, these observations are suggestive of behavioral interference, including the competitive displacement of pied tamarin by red-handed tamarins

Primary seed dispersal by three Neotropical seed-predating primates (Cacajao melanocephalus ouakary, Chiropotes chiropotes and Chiropotes albinasus)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)The Neotropics house two guilds of large arboreal vertebrate seed predators: parrots and the pitheciin primates. Both have diets dominated by immature fruits. The possibility of members of the Pitheciinae (genera Cacajao, Chiropotes and Pithecia) acting as occasional seed dispersers has been mooted, but not experimentally shown. We combined primate behavioural data and seed germination data from three separate field studies in the Brazilian states of Amazonas and Para to analyse patterns of post-consumption seed survivorship for seeds discarded by three pitheciin species (Cacajao melanocephalus ouakary, Chiropotes chiropotes and Chiropotes albinasus). We then calculated the frequency of dispersal events for four species eaten by C. m. ouakary. All three primate species dropped intact seeds while feeding, and 30.7% of 674 dropped seeds germinated ex situ. Undamaged seeds from unripe and ripe samples germinated (29.3% and 42.7%, respectively), and all three primate species carried some fruits up to 20 m from the parent tree before consuming them. Potential seed-dispersal events varied from 1 (Macrolobium acaciifolium) per fruiting cycle to more than 6500 (Duroia velutina), suggesting that there are differences in dispersal potential. In summary, although they are highly specialized seed predators, these primates may also act as important dispersers for some plant species, and effective dispersal is not restricted to ripe fruits, as immature fruits removed from a tree may continue to mature and the seeds later germinate, a much-neglected aspect of dispersal ecology. The possibility that similar events occur in parrots should be experimentally investigated.o TEXTO COMPLETO DESTE ARTIGO, ESTARÁ DISPONÍVEL À PARTIR DE AGOSTO DE 2015.286543555American Society of PrimatologistsColumbus Zoo Conservation FundLSB Leakey FoundationLeakey Foundation (UK)Linnean Society (Percy Sladen Memorial Fund)Margot Marsh Conservation FoundationMIL FoundationPittsburgh Zoo Conservation FundPrimate Conservation Inc.Roehampton UniversitySophie Danforth FundWildlife Conservation SocietyBDFFPSmithsonian Tropical Research InstituteArizona State UniversityFulbrightFundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Fundacao O Boticario de Protecao a NaturezaPrimate Action FundConselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)CNPq [BEV

A proposal for the common names for species of Chiropotes (Pitheciinae: Primates)

The common English name for the genus Chiropotes is currently bearded saki. We propose the use of 'cuxiu' as the common name for Chiropotes species, arguing that this term not only has deeper cultural and historical roots, but would mesh with the common name currently in use over the vast majority of the genus range. Cuxiu (pronounced 'coosh-e-oo') would be phylogenetically and taxonomically more appropriate, and less ambiguous, than the currently used term, and remove the implied close affiliation between Pithecia and Chiropotes. Finally, as an indigenously-derived name, it would fit with the common names in use for the other two genera in the sub-family Pitheciinae (uacari, Cacajao; saki, Pithecia), both of which also have indigenous origins.3507798