Marsupial position on life-history continua and the potential contribution of life-history traits to population growth.

Previous studies have suggested that mammal life history varies along the fast-slow continuum and that, in eutherians, this continuum is linked to variation in the potential contribution of survival and reproduction to population growth rate (λ). Fast eutherians mature early, have large litters and short lifespans, and exhibit high potential contribution of age at first reproduction and fertility to λ, while slow eutherians show high potential contribution of survival to λ. However, marsupials have typically been overlooked in comparative tests of mammalian life-history evolution. Here, we tested whether the eutherian life-history pattern extends to marsupials, and show that marsupial life-history trade-offs are organized along two major axes: (i) the reproductive output and dispersion axis, and (ii) the fast-slow continuum, with an additional association between adult survival and body mass. Life-history traits that potentially drive changes in λ are similar in eutherians and marsupials with slow life histories, but differ in fast marsupials; age at first reproduction is the most important trait contributing to λ and fertility contributes little. Marsupials have slower life histories than eutherians, and differences between these clades may derive from their contrasting reproductive modes; marsupials have slower development, growth and metabolism than eutherians of equivalent size.

Random placement models explain species richness and dissimilarity of frog assemblages within Atlantic Forest fragments.

Understanding the effects of random versus niche-based processes on biodiversity patterns is a central theme in ecology, and an important tool for predicting effects of habitat loss and fragmentation on biodiversity. We investigated the predictive power of random processes to explain species richness and species dissimilarity of amphibian assemblages in a fragmented tropical landscape of the Atlantic Forest of South America. We analyzed a large database of amphibian abundance and occupancy, sampled in 21 forest fragments ranging in size from 1.9 to 619 ha. We compared observed species richness and species dissimilarity with the outcomes of two null (random placement) models: 1- the traditional Coleman's area-based model and 2- an abundance-based model (based on the number of individuals observed in each fragment). We applied these models for all species combined, and separately for forest-dependent and habitat-generalist species. The abundance-based model fitted the observed species richness data better than the area-based model for all species, forest-dependent species, and generalist species. The area-based and the abundance-based models were also able to significantly explain species dissimilarity for all species and for generalists, but not for forest dependent species. The traditional area-based model assigned too many individuals to large fragments, thus failing to accurately explain species richness within patches across the landscape. Although niche-based processes may be important to structuring the regional pool of species in fragmented landscapes, our results suggest that part of the variation in species richness and species dissimilarity can be successfully explained by random placement models, especially for generalist species. Evaluating which factors cause variation in the number of individuals among patches should be a focus in future studies aiming to understand biodiversity patterns in fragmented landscapes.

Compreender os efeitos de processos aleatórios versus processos baseados em nicho nos padrões de biodiversidade é um tema central em ecologia e uma ferramenta importante para prever os efeitos da perda e fragmentação de habitat na biodiversidade. Nós investigamos o poder preditivo de processos aleatórios para explicar a riqueza e a dissimilaridade de espécies de assembleias de anfíbios em uma paisagem fragmentada tropical da Mata Atlântica da América do Sul. Analisamos um grande conjunto de dados de abundância e ocupação de anfíbios, amostrados em 21 fragmentos florestais com tamanhos de 1.9 a 619 ha. Comparamos a riqueza e a dissimilaridade de espécies observadas com os resultados de dois modelos nulos (posicionamento aleatório): 1- o modelo tradicional baseado em área de Coleman e 2 - um modelo baseado em abundância (com base no número de indivíduos observados em cada fragmento). Aplicamos esses modelos para todas as espécies combinadas e separadamente para espécies dependentes de floresta e espécies generalistas de habitat. O modelo baseado em abundância ajustou-se melhor aos dados observados de riqueza de espécies do que o modelo baseado em área para todas as espécies, espécies dependentes de floresta e espécies generalistas. Os modelos baseados em área e em abundância também foram capazes de explicar significativamente a dissimilaridade de espécies para todas as espécies e para generalistas, mas não para espécies dependentes de floresta. O modelo tradicional baseado em área atribuiu muitos indivíduos a grandes fragmentos, falhando assim em explicar com precisão a riqueza de espécies dentro de manchas na paisagem. Embora processos baseados em nicho possam ser importantes para estruturar o conjunto regional de espécies em paisagens fragmentadas, nossos resultados sugerem que parte da variação na riqueza e dissimilaridade de espécies pode ser explicada com sucesso por modelos de posicionamento aleatório, especialmente para espécies generalistas. Avaliar quais fatores causam variação no número de indivíduos entre manchas deve ser um foco em estudos futuros que visem compreender os padrões de biodiversidade em paisagens fragmentadas.

Sustainability issues in a tropical mega trail.

Sustainability is a target that involves many socio-ecological questions, depends on opportunities and combines different initiatives. This can be especially difficult in regions with high biodiversity scores, mega cities, high level of human populations and an intense and long-standing land use. Here, we show how a mega trail, named Atlantic Forest Trail, within the Brazilian Atlantic Forest can join the protection of biodiversity and sustainable tourism through a 4270 km corridor connecting protected areas and crossing a variety of landscapes. Further, we show some initiatives of ongoing biodiversity monitoring, and an analysis of ecological restoration in private lands that can be applied in many regions to improve habitat connectivity for both biodiversity and human use.

Small mammal responses to Amazonian forest islands are modulated by their forest dependence.

Hydroelectric dams have induced widespread loss, fragmentation and degradation of terrestrial habitats in lowland tropical forests. Yet their ecological impacts have been widely neglected, particularly in developing countries, which are currently earmarked for exponential hydropower development. Here we assess small mammal assemblage responses to Amazonian forest habitat insularization induced by the 28-year-old Balbina Hydroelectric Dam. We sampled small mammals on 25 forest islands (0.83-1466 ha) and four continuous forest sites in the mainland to assess the overall community structure and species-specific responses to forest insularization. We classified all species according to their degree of forest-dependency using a multi-scale approach, considering landscape, patch and local habitat characteristics. Based on 65,520 trap-nights, we recorded 884 individuals of at least 22 small mammal species. Species richness was best predicted by island area and isolation, with small islands (< 15 ha) harbouring an impoverished nested subset of species (mean ± SD: 2.6 ± 1.3 species), whereas large islands (> 200 ha; 10.8 ± 1.3 species) and continuous forest sites (∞ ha; 12.5 ± 2.5 species) exhibited similarly high species richness. Forest-dependent species showed higher local extinction rates and were often either absent or persisted at low abundances on small islands, where non-forest-dependent species became hyper-abundant. Species capacity to use non-forest habitat matrices appears to dictate small mammal success in small isolated islands. We suggest that ecosystem functioning may be highly disrupted on small islands, which account for 62.7% of all 3546 islands in the Balbina Reservoir.

Seasonal dynamics with compensatory effects regulate populations of tropical forest marsupials: a 16-year study.

The conditions that a population experiences during one season can affect the strength of density dependence in the following season. In the tropics, many populations face their biggest challenges in the dry season due to limited food and cold-dry conditions. Seasonal environmental changes can be especially problematic for small, short-lived, seasonally breeding endotherms. To investigate the effects of seasonality on population dynamics, we studied five marsupial species in the Brazilian Atlantic Forest, using a 16-year dataset. We tested if (1) compensatory density feedback is stronger in the dry season, due to the high population sizes and limited food; (2) lower temperatures and the overall abundance of small mammals negatively affect dry season population growth rates; and (3) rainfall, a proxy for food availability, is positively related to wet season population growth rates. Population growth rates were regressed against seasonal population sizes and exogenous variables, and analyzed with linear autoregressive models. Seasonal compensatory density feedback occurred in both seasons, with compensation processes in just one season being sufficient to allow population persistence. Rainfall and the overall abundance of small mammals had little influence on populations, while colder temperatures decreased population growth rate of smaller species in both seasons. Although the study marsupials share similar life histories and phylogeny, they varied with respect to the season when compensatory density feedback was strongest. Our results demonstrate that seasonality plays a key role in driving marsupial population dynamics, and highlight the need to account for seasonality in demographic studies even in tropical environments.

Edge effects and geometric constraints: a landscape-level empirical test.

Edge effects are pervasive in landscapes yet their causal mechanisms are still poorly understood. Traditionally, edge effects have been attributed to differences in habitat quality along the edge-interior gradient of habitat patches, under the assumption that no edge effects would occur if habitat quality was uniform. This assumption was questioned recently after the recognition that geometric constraints tend to reduce population abundances near the edges of habitat patches, the so-called geometric edge effect (GEE). Here, we present the first empirical, landscape-level evaluation of the importance of the GEE in shaping abundance patterns in fragmented landscapes. Using a data set on the distribution of small mammals across 18 forest fragments, we assessed whether the incorporation of the GEE into the analysis changes the interpretation of edge effects and the degree to which predictions based on the GEE match observed responses. Quantitative predictions were generated for each fragment using simulations that took into account home range, density and matrix use for each species. The incorporation of the GEE into the analysis changed substantially the interpretation of overall observed edge responses at the landscape scale. Observed abundances alone would lead to the conclusion that the small mammals as a group have no consistent preference for forest edges or interiors and that the black-eared opossum Didelphis aurita (a numerically dominant species in the community) has on average a preference for forest interiors. In contrast, incorporation of the GEE suggested that the small mammal community as a whole has a preference for forest edges, whereas D. aurita has no preference for forest edges or interiors. Unexplained variance in edge responses was reduced by the incorporation of GEE, but remained large, varying greatly on a fragment-by-fragment basis. This study demonstrates how to model and incorporate the GEE in analyses of edge effects and that this incorporation is necessary to properly interpret edge effects in landscapes. It also suggests that geometric constraints alone are unlikely to explain the variability in edge responses of a same species among different areas, highlighting the need to incorporate other ecological factors into explanatory models of edge effects.

Movement behaviour within and beyond perceptual ranges in three small mammals: effects of matrix type and body mass.

1. For animal species inhabiting heterogeneous landscapes, the tortuosity of the dispersal path is a key determinant of the success in locating habitat patches. Path tortuosity within and beyond perceptual range must differ, and may be differently affected by intrinsic attributes of individuals and extrinsic environmental factors. Understanding how these factors interact to determine path tortuosity allows more accurate inference of successful movements between habitat patches. 2. We experimentally determined the effects of intrinsic (body mass and species identity) and extrinsic factors (distance to nearest forest fragment and matrix type) on the tortuosity of movements of three forest-dwelling didelphid marsupials, in a fragmented landscape of the Atlantic Forest, Brazil. 3. A total of 202 individuals were captured in forest fragments and released in three unsuitable matrix types (mowed pasture, abandoned pasture and manioc plantation), carrying spool-and-line devices. 4. Twenty-four models were formulated representing a priori hypotheses of major determinants of path tortuosity, grouped in three scenarios (only intrinsic factors, only extrinsic factors and models with combinations of both), and compared using a model selection approach. Models were tested separately for individuals released within the perceptual range of the species, and for individuals released beyond the perceptual range. 5. Matrix type strongly affected path tortuosity, with more obstructed matrix types hampering displacement of animals. Body mass was more important than species identity to determine path tortuosity, with larger animals moving more linearly. Increased distance to the fragment resulted in more tortuous paths, but actually reflects a threshold in perceptual range: linear paths within perceptual range, tortuous paths beyond. 6. The variables tested explained successfully path tortuosity, but only for animals released within the perceptual range. Other factors, such as wind intensity and direction of plantation rows, may be more important for individuals beyond their perceptual range. 7. Simplistic scenarios considering only intrinsic or extrinsic factors are inadequate to predict path tortuosity, and to infer dispersal success in heterogeneous landscapes. Perceptual range represents a fundamental threshold where the effects of matrix type, body mass and individual behaviour change drastically.

Body size and extinction risk in Brazilian carnivores / Tamaño corporal como factor de amenaza en carnívoros brasileros

Because extinctions are not random across taxa, it is important for conservation biologists to identify the traits that make some species more vulnerable. Factors associated with vulnerability include small geographical ranges, low densities, high trophic level, "slow" life histories, body size, and tolerance to altered habitats. In this study we examined the relationship of body size, reproductive output, longevity, and extinction risk for carnivores occurring in Brazil. We used generalized linear models analyses on phylogenetically independent contrasts to test the effect of body size alone, and the combined effect of body size, litter size and longevity on extinction risk. Body size appeared in the two best models according to the selection criteria (AIC), and it was the most plausible bionomic variable associated with extinction risk. Litter size and longevity, bionomic traits previously associated with threat risk of Brazilian carnivores, were implausible. The higher extinction risk for larger species could result from body size influencing vulnerability to different human activities, such as killing, habitat destruction and fragmentation, and the small size of natural reserves.

Debido a que las extinciones no son aleatorias a través de los diferentes taxa, es importante para los biólogos de la conservación identificar las características que hacen a algunas especies más vulnerables. Los factores asociados con la vulnerabilidad incluyen distribuciones geográficas pequeñas, densidades poblacionales bajas, niveles tróficos altos, historias de vida "lentas", tamaño del cuerpo, y tolerancia a habitats alterados. En este estudio examinamos la relación del tamaño corporal, el potencial reproductivo y la longevidad con el riesgo de extinción para los carnívoros brasileros. Realizamos análisis de modelos lineares generalizados con contrastes filogenéticamente independientes para probar el efecto del tamaño corporal solo, y el efecto combinado del tamaño corporal, el tamaño de la camada y la longevidad sobre el riesgo de extinción. El tamaño del cuerpo apareció en los dos mejores modelos de acuerdo con el criterio de selección (AIC) y es la variable bionómica más plausible afectando el riesgo de extinción. El tamaño de la prole y longevidad, otras variables bionómicas que han presentado tal efecto en otros estudios, fueron implausibles. La mayor probabilidad de amenaza para las especies grandes puede deberse a que el tamaño del cuerpo afecta la vulnerabilidad a diferentes actividades humanas como la caza, la destrucción y fragmentación del hábitat, y al tamaño reducido de la mayoría de áreas protegidas.

Apparent competition through facilitation between Melanoides tuberculata and Biomphalaria glabrata and the control of schistosomiasis.

Interactions between two species that result in reduced growth rates for both and extinction of one of the species are generally considered cases of asymmetric interspecific competition. Exploitative or interference competition is the usual mechanism invoked. Here we describe another mechanism producing the same result, named apparent competition through facilitation (ACF), observed between Melanoides tuberculata and Biomphalaria glabrata populations. The superior competitor actually gives some benefit to the other species, whose population becomes unstable with progressively increasing oscillations, leading to extinction. A model of ACF using difference equations suggests initial dynamics distinct from traditional interspecific competition. The dynamics of two freshwater snails in the field and in laboratory experiments suggest ACF, and these relations should be considered in studies of schistosomiasis control. ACF could occur in natural populations, but might have gone undetected because the final result is similar to traditional interspecific competition.

Apparent competition through facilitation between Melanoides tuberculata and Biomphalaria glabrata and the control of schistosomiasis

Interactions between two species that result in reduced growth rates for both and extinction of one of the species are generally considered cases of asymmetric interspecific competition. Exploitative or interference competition is the usual mechanism invoked. Here we describe another mechanism producing the same result, named apparent competition through facilitation (ACF), observed between Melanoides tuberculata and Biomphalaria glabrata populations. The superior competitor actually gives some benefit to the other species, whose population becomes unstable with progressively increasing oscillations, leading to extinction. A model of ACF using difference equations suggests initial dynamics distinct from traditional interspecific competition. The dynamics of two freshwater snails in the field and in laboratory experiments suggest ACF, and these relations should be considered in studies of schistosomiasis control. ACF could occur in natural populations, but might have gone undetected because the final result is similar to traditional interspecific competition

Interaction between the intermediate host of Schistosomiasis in Brazil Biomphalaria glabrata (Planorbidae) and a possible competitor Melanoides tuberculata (Thiaridae): I. Laboratory experiments.

The biological control of Biomphalaria glabrata, intermediate host of Schistosoma mansoni, is one the accepted options to fight schistosomiasis. One of the most promising candidates to control B. glabrata is the snail Melanoides tuberculata, a potential competitor. However, the mechanisms of interaction between the two species are not clear. Our objective is to determine if M. tuberculata indeed compete with B. glabrata, using two laboratory experiments. In Experiment 1, we tested the effect of the presence of M. tuberculata on the fecundity and mortality rates of B. glabrata. In Experiment 2, we tested if there was a direct or indirect interaction between the two species. In Experiment 1, M. tuberculata was eliminated after the peak in reproductive activity of B. glabrata. In Experiment 2, B. glabrata produced more egg masses when raised with M. tuberculata. The conditions leading to this unexpected positive effect of M. tuberculata on the fecundity of B. glabrata need further clarification, but emphasize that detailed studies of the interaction between these species in the conditions of the local environment should be considered.

Interaction between the Intermediate Host of Schistosomiasis in Brazil Biomphalaria glabrata (Planorbidae) and a Possible Competitor Melanoides tuberculata (Thiaridae): I. Laboratory Experiments

The biological control of Biomphalaria glabrata, intermediate host of Schistosoma mansoni, is one the accepted options to fight schistosomiasis. One of the most promising candidates to control B. glabrata is the snail Melanoides tuberculata, a potential competitor. However, the mechanisms of interaction between the two species are not clear. Our objective is to determine if M. tuberculata indeed compete with B. glabrata, using two laboratory experiments. In Experiment 1, we tested the effect of the presence of M. tuberculata on the fecundity and mortality rates of B. glabrata. In Experiment 2, we tested if there was a direct or indirect interaction between the two species. In Experiment 1, M. tuberculata was eliminated after the peak in reproductive activity of B. glabrata. In Experiment 2, B. glabrata produced more egg masses when raised with M. tuberculata. The conditions leading to this unexpected positive effect of M. tuberculata on the fecundity of B. glabrata need further clarification, but emphasize that detailed studies of the interaction between these species in the conditions of the local environment should be considered