Smarter foragers do not forage smarter: a test of the diet hypothesis for brain expansion.

A leading hypothesis for the evolution of large brains in humans and other species is that a feedback loop exists whereby intelligent animals forage more efficiently, which results in increased energy intake that fuels the growth and maintenance of large brains. We test this hypothesis for the first time with high-resolution tracking data from four sympatric, frugivorous rainforest mammal species (42 individuals) and drone-based maps of their predominant feeding trees. We found no evidence that larger-brained primates had more efficient foraging paths than smaller brained procyonids. This refutes a key assumption of the fruit-diet hypothesis for brain evolution, suggesting that other factors such as temporal cognition, extractive foraging or sociality have been more important for brain evolution.

Invasive wild deer exhibit environmental niche shifts in Australia: Where to from here?

Invasive species have established populations around the world and, in the process, characteristics of their realized environmental niches have changed. Because of their popularity as a source of game, deer have been introduced to, and become invasive in, many different environments around the world. As such, deer should provide a good model system in which to test environmental niche shifts. Using the current distributions of the six deer species present in Australia, we quantified shifts in their environmental niches that occurred since introduction; we determined the differences in suitable habitat between their international (native and invaded) and their Australian ranges. Given knowledge of their Australian habitat use, we then modeled the present distribution of deer in Australia to assess habitat suitability, in an attempt to predict future deer distributions. We show that the Australian niches of hog (Axis porcinus), fallow (Dama dama), red (Cervus elaphus), rusa (C. timorensis), and sambar deer (C. unicolor), but not chital deer (A. axis), were different to their international ranges. When we quantified the potential range of these six species in Australia, chital, hog, and rusa deer had the largest areas of suitable habitat outside their presently occupied habitat. The other three species had already expanded outside the ranges that we predicted as suitable. Here, we demonstrate that deer have undergone significant environmental niche shifts following introduction into Australia, and these shifts are important for predicting the future spread of these invasive species. It is important to note that current Australian and international environmental niches did not necessarily predict range expansions, thus wildlife managers should treat these analyses as conservative estimates.

Dancing to a different tune: changing reproductive seasonality in an introduced chital deer population.

Male and female reproductive behaviour is typically synchronised. In species such as those in the family Cervidae, reproductive timing is often cued by photoperiod, although in females, it can be dependent on body condition. When a species is introduced to a novel environment, the environment changes, or responses of the sexes to such cues differ, asynchronous reproductive behaviour between males and females may occur. We investigated the seasonality of reproductive behaviour in introduced chital deer in northern Queensland by examining male antler phase in relation to female conception rates. We then analysed the influence of different variables likely to affect the timing of male and female reproductive physiology. The lowest percentage of chital in hard antler in any 1 month in this study was 35% (Fig. 1), but the average value was closer to 50%, thus there was a seasonal peak in antler phase linked with photoperiod. Females conceived at any time of year, but were strongly influenced by the amount of rainfall 3 months prior to conception. This resulted in varying conception peaks year-to-year that often did not correspond to the male's peak in hard antler. In this system, a proportion of males and females were physiologically and behaviourally ready to mate at any time of the year. We predict that differences in the timing of the peaks between the males and females will lead to increased reproductive skew (variation in reproductive success among individual males). This pattern may select for different mating strategies or physiological mechanisms to increase reproductive success. Fig. 1 The average percentage of male chital deer in hard antler by month from 2014 to 2019 in north Queensland. Values above the bars indicate the total number of males that were sampled in each month and the error bars indicate the standard error. In the month with the lowest % males in hard antler in the entire study (November, 2017), 35% of males were in hard antler.

Laying low: Rugged lowland rainforest preferred by feral cats in the Australian Wet Tropics.

Invasive mesopredators are responsible for the decline of many species of native mammals worldwide. Feral cats have been causally linked to multiple extinctions of Australian mammals since European colonization. While feral cats are found throughout Australia, most research has been undertaken in arid habitats, thus there is a limited understanding of feral cat distribution, abundance, and ecology in Australian tropical rainforests. We carried out camera-trapping surveys at 108 locations across seven study sites, spanning 200 km in the Australian Wet Tropics. Single-species occupancy analysis was implemented to investigate how environmental factors influence feral cat distribution. Feral cats were detected at a rate of 5.09 photographs/100 days, 11 times higher than previously recorded in the Australian Wet Tropics. The main environmental factors influencing feral cat occupancy were a positive association with terrain ruggedness, a negative association with elevation, and a higher affinity for rainforest than eucalypt forest. These findings were consistent with other studies on feral cat ecology but differed from similar surveys in Australia. Increasingly harsh and consistently wet weather conditions at higher elevations, and improved shelter in topographically complex habitats may drive cat preference for lowland rainforest. Feral cats were positively associated with roads, supporting the theory that roads facilitate access and colonization of feral cats within more remote parts of the rainforest. Higher elevation rainforests with no roads could act as refugia for native prey species within the critical weight range. Regular monitoring of existing roads should be implemented to monitor feral cats, and new linear infrastructure should be limited to prevent encroachment into these areas. This is pertinent as climate change modeling suggests that habitats at higher elevations will become similar to lower elevations, potentially making the environment more suitable for feral cat populations.

Effects of body size on estimation of mammalian area requirements.

Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home-range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied block cross-validation to quantify bias in empirical home-range estimates. Area requirements of mammals <10 kg were underestimated by a mean approximately15%, and species weighing approximately100 kg were underestimated by approximately50% on average. Thus, we found area estimation was subject to autocorrelation-induced bias that was worse for large species. Combined with the fact that extinction risk increases as body mass increases, the allometric scaling of bias we observed suggests the most threatened species are also likely to be those with the least accurate home-range estimates. As a correction, we tested whether data thinning or autocorrelation-informed home-range estimation minimized the scaling effect of autocorrelation on area estimates. Data thinning required an approximately93% data loss to achieve statistical independence with 95% confidence and was, therefore, not a viable solution. In contrast, autocorrelation-informed home-range estimation resulted in consistently accurate estimates irrespective of mass. When relating body mass to home range size, we detected that correcting for autocorrelation resulted in a scaling exponent significantly >1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum.

Efectos del Tamaño Corporal sobre la Estimación de los Requerimientos de Área de Mamíferos Resumen La cuantificación precisa de los requerimientos de área de una especie es un prerrequisito para que la conservación basada en áreas sea efectiva. Esto comúnmente implica la recolección de datos de rastreo de la especie de interés para después realizar análisis de la distribución local. De manera problemática, la autocorrelación en los datos de rastreo puede resultar en una subestimación grave de las necesidades de espacio. Con base en trabajos previos, formulamos una hipótesis en la que supusimos que la magnitud de la subestimación varía con la masa corporal, una relación que podría tener implicaciones serias para la conservación. Para probar esta hipótesis en mamíferos terrestres, estimamos las áreas de distribución local con las ubicaciones en GPS de 757 individuos de 61 especies de mamíferos distribuidas mundialmente con una masa corporal entre 0.4 y 4,000 kg. Después aplicamos una validación cruzada en bloque para cuantificar el sesgo en estimaciones empíricas de la distribución local. Los requerimientos de área de los mamíferos <10 kg fueron subestimados por una media ∼15% y las especies con una masa ∼100 kg fueron subestimadas en ∼50% en promedio. Por lo tanto, encontramos que la estimación del área estaba sujeta al sesgo inducido por la autocorrelación, el cual era peor para las especies de talla grande. En combinación con el hecho de que el riesgo de extinción incrementa conforme aumenta la masa corporal, el escalamiento alométrico del sesgo que observamos sugiere que la mayoría de las especies amenazadas también tienen la probabilidad de ser aquellas especies con las estimaciones de distribución local menos acertadas. Como corrección, probamos si la reducción de datos o la estimación de la distribución local informada por la autocorrelación minimizan el efecto de escalamiento que tiene la autocorrelación sobre las estimaciones de área. La reducción de datos requirió una pérdida de datos del ∼93% para lograr la independencia estadística con un 95% de confianza y por lo tanto no fue una solución viable. Al contrario, la estimación de la distribución local informada por la autocorrelación resultó en estimaciones constantemente precisas sin importar la masa corporal. Cuando relacionamos la masa corporal con el tamaño de la distribución local, detectamos que la corrección de la autocorrelación resultó en un exponente de escalamiento significativamente >1, lo que significa que el escalamiento de la relación cambió sustancialmente en el extremo superior del espectro de la masa corporal.

Phylogeographic and diversification patterns of the white-nosed coati (Nasua narica): Evidence for south-to-north colonization of North America.

White-nosed coatis (Nasua narica) are widely distributed throughout North, Central, and South America, but the patterns of temporal and spatial diversification that have contributed to this distribution are unknown. In addition, the biogeographic history of procyonid species in the Americas remains contentious. Using sequences from three mitochondrial loci (Cytochrome b, NAHD5 and 16S rRNA; 2201 bp) and genotypes from 11 microsatellite loci, we analyzed genetic diversity to determine phylogeographic patterns, genetic structure, divergence times, and gene flow among Nasua narica populations throughout the majority of the species' range. We also estimated the ancestral geographic range of N. narica and other procyonid species. We found a high degree of genetic structure and divergence among populations that conform to five evolutionarily significant units. The most southerly distributed population (Panama) branched off much earlier (∼3.8 million years ago) than the northern populations (<1.2 million years ago). Estimated gene flow among populations was low and mostly northwards and westwards. The phylogeographic patterns within N. narica are associated with geographic barriers and habitat shifts likely caused by Pliocene-Pleistocene climate oscillations. Significantly, our findings suggest the dispersal of N. narica was south-to-north beginning in the Pliocene, not in the opposite direction during the Pleistocene as suggested by the fossil record, and that the most recent common ancestor for coati species was most likely distributed in South or Central America six million years ago. Our study implies the possibility that the diversification of Nasua species, and other extant procyonid lineages, may have occurred in South America.

Raccoon contact networks predict seasonal susceptibility to rabies outbreaks and limitations of vaccination.

Infectious disease transmission often depends on the contact structure of host populations. Although it is often challenging to capture the contact structure in wild animals, new technology has enabled biologists to obtain detailed temporal information on wildlife social contacts. In this study, we investigated the effects of raccoon contact patterns on rabies spread using network modelling. Raccoons (Procyon lotor) play an important role in the maintenance of rabies in the United States. It is crucial to understand how contact patterns influence the spread of rabies in raccoon populations in order to design effective control measures and to prevent transmission to human populations and other animals. We constructed a dynamic system of contact networks based on empirical data from proximity logging collars on a wild suburban raccoon population and then simulated rabies spread across these networks. Our contact networks incorporated the number and duration of raccoon interactions. We included differences in contacts according to sex and season, and both short-term acquaintances and long-term associations. Raccoons may display different behaviours when infectious, including aggression (furious behaviour) and impaired mobility (dumb behaviour); the network model was used to assess the impact of potential behavioural changes in rabid raccoons. We also tested the effectiveness of different vaccination coverage levels. Our results demonstrate that when rabies enters a suburban raccoon population, the likelihood of a disease outbreak affecting the majority of the population is high. Both the magnitude of rabies outbreaks and the speed of rabies spread depend strongly on the time of year that rabies is introduced into the population. When there is a combination of dumb and furious behaviours in the rabid raccoon population, there are similar outbreak sizes and speed of spread to when there are no behavioural changes due to rabies infection. By incorporating detailed data describing the variation in raccoon contact rates into a network modelling approach, we were able to show that suburban raccoon populations are highly susceptible to rabies outbreaks, that the risk of large outbreaks varies seasonally and that current vaccination target levels may be inadequate to prevent the spread of rabies within these populations. Our findings provide new insights into rabies dynamics in raccoon populations and have important implications for disease control.

Patterns of latrine use by raccoons (Procyon lotor) and implication for Baylisascaris procyonis transmission.

Mammals often use latrine sites for defecation, yet little is known about patterns of latrine use in many common species such as raccoons (Procyon lotor). Because raccoon latrines are important foci for the transmission of raccoon roundworm (Baylisascaris procyonis), documenting metrics of raccoon latrine use may have public health implications. Although some studies have provided evidence that multiple raccoons visit single latrine sites, exact latrine visitation patterns of raccoons have never been documented. We monitored raccoon latrine usage using proximity-logging collars placed at 15 latrine sites. We found that latrine sites were visited by multiple raccoons (range 1-7), and raccoons visited as many as six latrines during a 2-wk period. No sex differences were found in the number of latrines visited or time spent during visits. We posit that the use of multiple latrine sites by raccoons may lead to the pattern that rates of B. procyonis infection at latrines are greater than infection rates found in individual raccoon fecal samples. This in turn could lead to greater transmission of B. procyonis to paratenic hosts. Our results support the conclusion that raccoon latrines can be major foci for the infection and spread of B. procyonis.

Raccoon social networks and the potential for disease transmission.

Raccoons are an important vector of rabies and other pathogens. The degree to which these pathogens can spread through a raccoon population should be closely linked to association rates between individual raccoons. Most studies of raccoon sociality have found patterns consistent with low levels of social connectivity within populations, thus the likelihood of direct pathogen transmission between raccoons is theoretically low. We used proximity detecting collars and social network metrics to calculate the degree of social connectivity in an urban raccoon population for purposes of estimating potential pathogen spread. In contrast to previous assumptions, raccoon social association networks were highly connected, and all individuals were connected to one large social network during 15 out of 18 months of study. However, these metrics may overestimate the potential for a pathogen to spread through a population, as many of the social connections were based on relatively short contact periods. To more closely reflect varying probabilities of pathogen spread, we censored the raccoon social networks based on the total amount of time spent in close proximity between two individuals per month. As this time criteria for censoring the social networks increased from one to thirty minutes, corresponding measures of network connectivity declined. These findings demonstrate that raccoon populations are much more tightly connected than would have been predicted based on previous studies, but also point out that additional research is needed to calculate more precise transmission probabilities by infected individuals, and determine how disease infection changes normal social behaviors.

Directed seed dispersal towards areas with low conspecific tree density by a scatter-hoarding rodent.

Scatter-hoarding animals spread out cached seeds to reduce density-dependent theft of their food reserves. This behaviour could lead to directed dispersal into areas with lower densities of conspecific trees, where seed and seedling survival are higher, and could profoundly affect the spatial structure of plant communities. We tested this hypothesis with Central American agoutis and Astrocaryum standleyanum palm seeds on Barro Colorado Island, Panama. We radio-tracked seeds as they were cached and re-cached by agoutis, calculated the density of adult Astrocaryum trees surrounding each cache, and tested whether the observed number of trees around seed caches declined more than expected under random dispersal. Seedling establishment success was negatively dependent on seed density, and agoutis carried seeds towards locations with lower conspecific tree densities, thus facilitating the escape of seeds from natural enemies. This behaviour may be a widespread mechanism leading to highly effective seed dispersal by scatter-hoarding animals.

Thieving rodents as substitute dispersers of megafaunal seeds.

The Neotropics have many plant species that seem to be adapted for seed dispersal by megafauna that went extinct in the late Pleistocene. Given the crucial importance of seed dispersal for plant persistence, it remains a mystery how these plants have survived more than 10,000 y without their mutualist dispersers. Here we present support for the hypothesis that secondary seed dispersal by scatter-hoarding rodents has facilitated the persistence of these large-seeded species. We used miniature radio transmitters to track the dispersal of reputedly megafaunal seeds by Central American agoutis, which scatter-hoard seeds in shallow caches in the soil throughout the forest. We found that seeds were initially cached at mostly short distances and then quickly dug up again. However, rather than eating the recovered seeds, agoutis continued to move and recache the seeds, up to 36 times. Agoutis dispersed an estimated 35% of seeds for >100 m. An estimated 14% of the cached seeds survived to the next year, when a new fruit crop became available to the rodents. Serial video-monitoring of cached seeds revealed that the stepwise dispersal was caused by agoutis repeatedly stealing and recaching each other's buried seeds. Although previous studies suggest that rodents are poor dispersers, we demonstrate that communities of rodents can in fact provide highly effective long-distance seed dispersal. Our findings suggest that thieving scatter-hoarding rodents could substitute for extinct megafaunal seed dispersers of tropical large-seeded trees.

Kinship shapes affiliative social networks but not aggression in ring-tailed coatis.

Animal groups typically contain individuals with varying degrees of genetic relatedness, and this variation in kinship has a major influence on patterns of aggression and affiliative behaviors. This link between kinship and social behavior underlies socioecological models which have been developed to explain how and why different types of animal societies evolve. We tested if kinship and age-sex class homophily in two groups of ring-tailed coatis (Nasua nasua) predicted the network structure of three different social behaviors: 1) association, 2) grooming, and 3) aggression. Each group was studied during two consecutive years, resulting in four group-years available for analysis (total of 65 individuals). Association patterns were heavily influenced by agonistic interactions which typically occurred during feeding competition. Grooming networks were shaped by mother-offspring bonds, female-female social relationships, and a strong social attraction to adult males. Mother-offspring pairs were more likely to associate and groom each other, but relatedness had no effect on patterns of aggressive behavior. Additionally, kinship had little to no effect on coalitionary support during agonistic interactions. Adult females commonly came to the aid of juveniles during fights with other group members, but females often supported juveniles who were not their offspring (57% of coalitionary interactions). These patterns did not conform to predictions from socioecological models.

Familiarity breeds progeny: sociality increases reproductive success in adult male ring-tailed coatis (Nasua nasua).

The ring-tailed coati (Nasua nasua) is the only coati species in which social groups contain an adult male year round, although most males live solitarily. We compared reproductive success of group living and solitary adult male coatis to determine the degree to which sociality affects reproductive success. Coati mating is highly seasonal and groups of female coatis come into oestrus during the same 1-2 week period. During the mating season, solitary adult males followed groups and fought with the group living male. This aggression was presumably to gain access to receptive females. We expected that high reproductive synchrony would make it difficult or impossible for the one group living male to monopolize and defend the group of oestrous females. However, we found that group living males sired between 67-91% of the offspring in their groups. This reproductive monopolization is much higher than other species of mammals with comparably short mating seasons. Clearly, living in a group greatly enhanced a male's reproductive success. At the same time, at least 50% of coati litters contained offspring sired by extra-group males (usually only one offspring per litter); thus, resident males could not prevent extra-group matings. The resident male's reproductive advantage may reflect female preference for a resident male strong enough to fend off competing males.

Costs and benefits of within-group spatial position: a feeding competition model.

An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.