Prey availability and temporal partitioning modulate felid coexistence in Neotropical forests

Carnivores have long been used as model organisms to examine mechanisms that allow coexistence among ecologically similar species. Interactions between carnivores, including competition and predation, comprise important processes regulating local community structure and diversity. We use data from an intensive camera-trapping monitoring program across eight Neotropical forest sites to describe the patterns of spatiotemporal organization of a guild of five sympatric cat species: jaguar (Panthera onca), puma (Puma concolor), ocelot (Leopardus pardalis), jaguarundi (Herpailurus yagouaroundi) and margay (Leopardus wiedii). For the three largest cat species, we developed multi-stage occupancy models accounting for habitat characteristics (landscape complexity and prey availability) and models accounting for species interactions (occupancy estimates of potential competitor cat species). Patterns of habitat-use were best explained by prey availability, rather than habitat structure or species interactions, with no evidence of negative associations of jaguar on puma and ocelot occupancy or puma on ocelot occupancy. We further explore temporal activity patterns and overlap of all five felid species. We observed a moderate temporal overlap between jaguar, puma and ocelot, with differences in their activity peaks, whereas higher temporal partitioning was observed between jaguarundi and both ocelot and margay. Lastly, we conducted temporal overlap analysis and calculated species activity levels across study sites to explore if shifts in daily activity within species can be explained by varying levels of local competition pressure. Activity patterns of ocelots, jaguarundis and margays were similarly bimodal across sites, but pumas exhibited irregular activity patterns, most likely as a response to jaguar activity. Activity levels were similar among sites and observed differences were unrelated to competition or intraguild killing risk. Our study reveals apparent spatial and temporal partitioning for most of the species pairs analyzed, with prey abundance being more important than species interactions in governing the local occurrence and spatial distribution of Neotropical forest felids

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

An empirical evaluation of camera trap study design: How many, how long and when?

Abstract Camera traps deployed in grids or stratified random designs are a well‐established survey tool for wildlife but there has been little evaluation of study design parameters. We used an empirical subsampling approach involving 2,225 camera deployments run at 41 study areas around the world to evaluate three aspects of camera trap study design (number of sites, duration and season of sampling) and their influence on the estimation of three ecological metrics (species richness, occupancy and detection rate) for mammals. We found that 25–35 camera sites were needed for precise estimates of species richness, depending on scale of the study. The precision of species‐level estimates of occupancy (ψ) was highly sensitive to occupancy level, with 0.75) species, but more than 150 camera sites likely needed for rare (ψ < 0.25) species. Species detection rates were more difficult to estimate precisely at the grid level due to spatial heterogeneity, presumably driven by unaccounted habitat variability factors within the study area. Running a camera at a site for 2 weeks was most efficient for detecting new species, but 3–4 weeks were needed for precise estimates of local detection rate, with no gains in precision observed after 1 month. Metrics for all mammal communities were sensitive to seasonality, with 37%–50% of the species at the sites we examined fluctuating significantly in their occupancy or detection rates over the year. This effect was more pronounced in temperate sites, where seasonally sensitive species varied in relative abundance by an average factor of 4–5, and some species were completely absent in one season due to hibernation or migration. We recommend the following guidelines to efficiently obtain precise estimates of species richness, occupancy and detection rates with camera trap arrays: run each camera for 3–5 weeks across 40–60 sites per array. We recommend comparisons of detection rates be model based and include local covariates to help account for small‐scale variation. Furthermore, comparisons across study areas or times must account for seasonality, which could have strong impacts on mammal communities in both tropical and temperate sites

Phylogenetic diversity of Amazonian tree communities

Aim: To examine variation in the phylogenetic diversity (PD) of tree communities across geographical and environmental gradients in Amazonia. Location: Two hundred and eighty-three c. 1 ha forest inventory plots from across Amazonia. Methods: We evaluated PD as the total phylogenetic branch length across species in each plot (PDss), the mean pairwise phylogenetic distance between species (MPD), the mean nearest taxon distance (MNTD) and their equivalents standardized for species richness (ses.PDss, ses.MPD, ses.MNTD). We compared PD of tree communities growing (1) on substrates of varying geological age; and (2) in environments with varying ecophysiological barriers to growth and survival. Results: PDss is strongly positively correlated with species richness (SR), whereas MNTD has a negative correlation. Communities on geologically young- and intermediate-aged substrates (western and central Amazonia respectively) have the highest SR, and therefore the highest PDss and the lowest MNTD. We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses.PDss and ses.MNTD. MPD and ses.MPD are strongly correlated with how evenly taxa are distributed among the three principal angiosperm clades and are both highest in western Amazonia. Meanwhile, seasonally dry tropical forest (SDTF) and forests on white sands have low PD, as evaluated by any metric. Main conclusions: High ses.PDss and ses.MNTD reflect greater lineage diversity in communities. We suggest that high ses.PDss and ses.MNTD in western Amazonia results from its favourable, easy-to-colonize environment, whereas high values in the Brazilian and Guianan Shields may be due to accumulation of lineages over a longer period of time. White-sand forests and SDTF are dominated by close relatives from fewer lineages, perhaps reflecting ecophysiological barriers that are difficult to surmount evolutionarily. Because MPD and ses.MPD do not reflect lineage diversity per se, we suggest that PDss, ses.PDss and ses.MNTD may be the most useful diversity metrics for setting large-scale conservation priorities

Phylogenetic diversity of Amazonian tree communities

This is the peer reviewed version of the following article: Honorio Coronado, E. N., Dexter, K. G., Pennington, R. T., Chave, J., Lewis, S. L., Alexiades, M. N., Alvarez, E., Alves de Oliveira, A., Amaral, I. L., Araujo-Murakami, A., Arets, E. J. M. M., Aymard, G. A., Baraloto, C., Bonal, D., Brienen, R., Cerón, C., Cornejo Valverde, F., Di Fiore, A., Farfan-Rios, W., Feldpausch, T. R., Higuchi, N., Huamantupa-Chuquimaco, I., Laurance, S. G., Laurance, W. F., López-Gonzalez, G., Marimon, B. S., Marimon-Junior, B. H., Monteagudo Mendoza, A., Neill, D., Palacios Cuenca, W., Peñuela Mora, M. C., Pitman, N. C. A., Prieto, A., Quesada, C. A., Ramirez Angulo, H., Rudas, A., Ruschel, A. R., Salinas Revilla, N., Salomão, R. P., Segalin de Andrade, A., Silman, M. R., Spironello, W., ter Steege, H., Terborgh, J., Toledo, M., Valenzuela Gamarra, L., Vieira, I. C. G., Vilanova Torre, E., Vos, V., Phillips, O. L. (2015), Phylogenetic diversity of Amazonian tree communities. Diversity and Distributions, 21: 1295–1307. doi: 10.1111/ddi.12357, which has been published in final form at 10.1111/ddi.12357Aim: To examine variation in the phylogenetic diversity (PD) of tree communities across geographical and environmental gradients in Amazonia. Location: Two hundred and eighty-three c. 1 ha forest inventory plots from across Amazonia. Methods: We evaluated PD as the total phylogenetic branch length across species in each plot (PDss), the mean pairwise phylogenetic distance between species (MPD), the mean nearest taxon distance (MNTD) and their equivalents standardized for species richness (ses.PDss, ses.MPD, ses.MNTD). We compared PD of tree communities growing (1) on substrates of varying geological age; and (2) in environments with varying ecophysiological barriers to growth and survival. Results: PDss is strongly positively correlated with species richness (SR), whereas MNTD has a negative correlation. Communities on geologically young- and intermediate-aged substrates (western and central Amazonia respectively) have the highest SR, and therefore the highest PDss and the lowest MNTD. We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses.PDss and ses.MNTD. MPD and ses.MPD are strongly correlated with how evenly taxa are distributed among the three principal angiosperm clades and are both highest in western Amazonia. Meanwhile, seasonally dry tropical forest (SDTF) and forests on white sands have low PD, as evaluated by any metric. Main conclusions: High ses.PDss and ses.MNTD reflect greater lineage diversity in communities. We suggest that high ses.PDss and ses.MNTD in western Amazonia results from its favourable, easy-to-colonize environment, whereas high values in the Brazilian and Guianan Shields may be due to accumulation of lineages over a longer period of time. White-sand forests and SDTF are dominated by close relatives from fewer lineages, perhaps reflecting ecophysiological barriers that are difficult to surmount evolutionarily. Because MPD and ses.MPD do not reflect lineage diversity per se, we suggest that PDss, ses.PDss and ses.MNTD may be the most useful diversity metrics for setting large-scale conservation priorities.FINCyT - PhD studentshipSchool of Geography of the University of LeedsRoyal Botanic Garden EdinburghNatural Environment Research Council (NERC)Gordon and Betty Moore FoundationEuropean Union's Seventh Framework ProgrammeERCCNPq/PELDNSF - Fellowshi

Fast demographic traits promote high diversification rates of Amazonian trees.

The Amazon rain forest sustains the world's highest tree diversity, but it remains unclear why some clades of trees are hyperdiverse, whereas others are not. Using dated phylogenies, estimates of current species richness and trait and demographic data from a large network of forest plots, we show that fast demographic traits ? short turnover times ? are associated with high diversification rates across 51 clades of canopy trees. This relationship is robust to assuming that diversification rates are either constant or decline over time, and occurs in a wide range of Neotropical tree lineages. This finding reveals the crucial role of intrinsic, ecological variation among clades for understanding the origin of the remarkable diversity of Amazonian trees and forests

Assessment of motor functioning in the preschool period

The assessment of motor functioning in young children has become increasingly important in recent years with the acknowledgement that motor impairment is linked with cognitive, language, social and emotional difficulties. However, there is no one gold standard assessment tool to investigate motor ability in children. The aim of the current paper was to discuss the issues related to the assessment of motor ability in young pre-school children and to provide guidelines on the best approach for motor assessment. The paper discusses the maturational changes in brain development at the preschool level in relation to motor ability. Other issues include sex differences in motor ability at this young age, and evidence for this in relation to sociological versus biological influences. From the previous literature it is unclear what needs to be assessed in relation to motor functioning. Should the focus be underlying motor processes or movement skill assessment? Several key assessment tools are discussed that produce a general measure of motor performance followed by a description of tools that assess specific skills, such as fine and gross motor, ball and graphomotor skills. The paper concludes with recommendations on the best approach in assessing motor function in pre-school children