Climate variability and aridity modulate the role of leaf shelters for arthropods: A global experiment.

Current climate change is disrupting biotic interactions and eroding biodiversity worldwide. However, species sensitive to aridity, high temperatures, and climate variability might find shelter in microclimatic refuges, such as leaf rolls built by arthropods. To explore how the importance of leaf shelters for terrestrial arthropods changes with latitude, elevation, and climate, we conducted a distributed experiment comparing arthropods in leaf rolls versus control leaves across 52 sites along an 11,790 km latitudinal gradient. We then probed the impact of short- versus long-term climatic impacts on roll use, by comparing the relative impact of conditions during the experiment versus average, baseline conditions at the site. Leaf shelters supported larger organisms and higher arthropod biomass and species diversity than non-rolled control leaves. However, the magnitude of the leaf rolls' effect differed between long- and short-term climate conditions, metrics (species richness, biomass, and body size), and trophic groups (predators vs. herbivores). The effect of leaf rolls on predator richness was influenced only by baseline climate, increasing in magnitude in regions experiencing increased long-term aridity, regardless of latitude, elevation, and weather during the experiment. This suggests that shelter use by predators may be innate, and thus, driven by natural selection. In contrast, the effect of leaf rolls on predator biomass and predator body size decreased with increasing temperature, and increased with increasing precipitation, respectively, during the experiment. The magnitude of shelter usage by herbivores increased with the abundance of predators and decreased with increasing temperature during the experiment. Taken together, these results highlight that leaf roll use may have both proximal and ultimate causes. Projected increases in climate variability and aridity are, therefore, likely to increase the importance of biotic refugia in mitigating the effects of climate change on species persistence.

Reconciling biome-wide conservation of an apex carnivore with land-use economics in the increasingly threatened Pantanal wetlands.

Conservation of carnivores involves finding solutions to minimize habitat loss and human-wildlife conflict. Understanding the nature of land-use economics can allow us to mitigate both threats. In the Pantanal, the two main economic activities are cattle ranching and ecotourism, each of which directly and indirectly affect the persistence of jaguars (Panthera onca). To understand how the geography of these economic activities is related to jaguar populations, we developed a jaguar distribution model (JDM), livestock density model, and ecotourism lodge density model for the Pantanal. Due to the recent wildfires within the Pantanal, we also assess the impact of burnt areas that are suitable for jaguars, cattle ranching, and tourism. Our JDM indicate that 64% of the Pantanal holds suitable habitat for jaguars. However, jaguar habitat suitability was positively correlated with ecotourism, but negatively correlated with areas most suitable for intensive cattle-ranching. This demonstrates a biome-wide scenario compatible with jaguar conservation. Of particular concern, recent wildfires overlap most suitable areas for jaguars. If wildfires become increasingly frequent, this would represent a serious threat to jaguars and many other wildlife populations. We emphasize the global importance of the Pantanal wetland ecoregion as a key stronghold for long-term jaguar conservation.

The influence of spatial sampling scales on ant-plant interaction network architecture.

Despite great interest in metrics to quantify the structure of ecological networks, the effects of sampling and scale remain poorly understood. In fact, one of the most challenging issues in ecology is how to define suitable scales (i.e., temporal or spatial) to accurately describe and understand ecological systems. Here, we sampled a series of ant-plant interaction networks in the southern Brazilian Amazon rainforest in order to determine whether the spatial sampling scale, from local to regional, affects our understanding of the structure of these networks. To this end, we recorded ant-plant interactions in adjacent 25 × 30 m subplots (local sampling scale) nested within twelve 250 × 30 m plots (regional sampling scale). Moreover, we combined adjacent or random subplots and plots in order to increase the spatial sampling scales at the local and regional levels. We then calculated commonly used binary and quantitative network-level metrics for both sampling scales (i.e., number of species and interactions, nestedness, specialization and modularity), all of which encompass a wide array of structural patterns in interaction networks. We observed increasing species and interactions across sampling scales, and while most network descriptors remained relatively constant at the local level, there was more variation at the regional scale. Among all metrics, specialization was most constant across different spatial sampling scales. Furthermore, we observed that adjacent assembly did not generate more variation in network descriptor values compared to random assembly. This finding indicates that the spatially aggregated distribution of species/individuals and abiotic conditions does not affect the organization of these interacting assemblages. Our results have a direct impact on our empirical and theoretical understanding of the ecological dynamics of species interactions by demonstrating that small spatial sampling scales should suffice to record some patterns commonly found in ant-plant interaction networks in a highly diverse tropical rainforest.

Postponing the production of ant domatia as a strategy promoting an escape from flooding in an Amazonian myrmecophyte.

Background and Aims: Even when adapted to flooding environments, the spatial distribution, growing strategies and anti-herbivore defences of plants face stressful conditions. Here we describe the effects of flooding on carbon allocation on growth, domatia and leaf production, and the herbivory on the myrmecophyte domatia-bearing Tococa coronata Benth. (Melastomataceae) growing along river banks in the Amazon region. Methods: In an area of 80 000 m2 of riparian forest along the Juruena River we actively searched for individuals of T. coronata. In each plant we evaluated the size of the plant when producing the first domatium and determined its best predictor: (1) plant total height; (2) size of plants above flood level; or (3) length of time each plant spent underwater. We also compared the herbivory, internode elongation, foliar asymmetry and specific leaf weight between T. coronata individuals growing above and below the maximum flooding level. The distance to the river and the height of the first domatium produced were compared between T. coronata and its sympatric congener, T. bulifera. Key Results: We found that T. coronata invests in rapid growth in the early ontogenetic stages through an elongation of internodes rather than in constitutive anti-herbivore defences to leaves or domatia to exceed the maximum flooding level. Consequently, its leaf herbivory was higher when compared with those produced above the flooding level. Individuals with leaves above flood levels produce coriaceous leaves and ant-domatias. Thus, flooding seems to trigger changes in growth strategies of the species. Furthermore, T. coronata occurs within the flood level, whereas its congener T. bullifera invariably occurs at sites unreachable by floods. Conclusion: Even in conditions of high stress, T. coronata presents both physiological and adaptive strategies that allow for colonization and establishment within flooded regions. These mechanisms involve an extreme trade-off of postponing adult plant characteristics to rapid growth to escape flooding while minimizing carbon allocation to defence.

Assemblage and functional categorization of dung beetles (Coleoptera: Scarabaeinae) from the Pantanal.

The Pantanal is one of the world's largest tropical wetland areas and harbors high mammal biomass. There is no formal list of dung beetle species, and studies on their functional roles have never being carried out in Pantanal. In this study, we identified dung beetle species occurring in the north Pantanal region (Poconé sub-region, Brazil) and studied their functional organization, by measuring morphological, behavioral and phenological traits. We collected 25,278 individuals belonging to 17 genera and 35 species. We identified eight functional groups in the habitat: Noturnal Telecoprids, Diurnal Telecoprids, Nesting Endocoprids, Small Nonrollers, Nocturnal Nester Paracoprids, Big Nesters Paracoprids, Non Nesters Paracoprids and Diurnal Nesters Paracoprids. The functional groups were defined mostly by two reproductive traits and two niche differentiation traits related to the use of fecal resources. This high diversification of both species and functional roles shows the importance of the group in a habitat with strong variation in availability of habitat and resources.

Individual-based ant-plant networks: diurnal-nocturnal structure and species-area relationship.

Despite the importance and increasing knowledge of ecological networks, sampling effort and intrapopulation variation has been widely overlooked. Using continuous daily sampling of ants visiting three plant species in the Brazilian Neotropical savanna, we evaluated for the first time the topological structure over 24 h and species-area relationships (based on the number of extrafloral nectaries available) in individual-based ant-plant networks. We observed that diurnal and nocturnal ant-plant networks exhibited the same pattern of interactions: a nested and non-modular pattern and an average level of network specialization. Despite the high similarity in the ants' composition between the two collection periods, ant species found in the central core of highly interacting species totally changed between diurnal and nocturnal sampling for all plant species. In other words, this "night-turnover" suggests that the ecological dynamics of these ant-plant interactions can be temporally partitioned (day and night) at a small spatial scale. Thus, it is possible that in some cases processes shaping mutualistic networks formed by protective ants and plants may be underestimated by diurnal sampling alone. Moreover, we did not observe any effect of the number of extrafloral nectaries on ant richness and their foraging on such plants in any of the studied ant-plant networks. We hypothesize that competitively superior ants could monopolize individual plants and allow the coexistence of only a few other ant species, however, other alternative hypotheses are also discussed. Thus, sampling period and species-area relationship produces basic information that increases our confidence in how individual-based ant-plant networks are structured, and the need to consider nocturnal records in ant-plant network sampling design so as to decrease inappropriate inferences.

The structure of ant-plant ecological networks: is abundance enough?

Knowledge of the mechanisms that shape biodiversity is essential to understand the ecological and evolutionary dynamics of interacting species. Recent studies posit that most of the organization of mutualistic networks is shaped by differences in species abundance among interacting species. In this study, we examined the mutualism involving plants with extrafloral nectaries and their associated ants. We show empirically that the difference in abundance among ants on vegetation partially explains the network structure of mutualistic interactions and that it is independent of ant species compositions: an ant species that is abundant usually interacts with more plant species. Moreover, nested networks are generated by simple variation in ant abundance on foliage. However, in ant-plant mutualistic networks, nestedness was higher than in networks describing the occurrence of ants on plants without a food resource. Additionally, the plant and ant species with the highest number of interactions within these networks interacted more among themselves than expected under the assumption of an abundance-based, random mixing of individuals. We hypothesize that the dominance of these ant species occurs because these ants are able to outcompete other ant species when feeding on extrafloral nectaries and because of the presence of ecophysiological adaptations to utilize liquid food.

The geographic distribution of parasite-induced fruit mimicry in Cephalotes atratus (Formicidae: Myrmicinae).

Parasite distributions fundamentally depend on the distributions of their hosts but may be more restricted than their hosts. Host-parasite symbioses tend to be spatially aggregated, and widely distributed host-parasite relationships are rare. Here, we combine field observations with published collection data to document the current known distribution of the nematode, Myrmeconema neotropicum, which infects the Neotropical canopy ant Cephalotes atratus. We report 6 new records from different Brazilian ecosystems, bringing the total number of independent observations of this interaction to 11. The broad distribution of these data points suggests that M. neotropicum infects C. atratus throughout its geographic range, although possible disturbance effects and specific habitat associations of the interaction remain unknown.

Asymmetric dispersal and colonization success of Amazonian plant-ants queens.

BACKGROUND: The dispersal ability of queens is central to understanding ant life-history evolution, and plays a fundamental role in ant population and community dynamics, the maintenance of genetic diversity, and the spread of invasive ants. In tropical ecosystems, species from over 40 genera of ants establish colonies in the stems, hollow thorns, or leaf pouches of specialized plants. However, little is known about the relative dispersal ability of queens competing for access to the same host plants. METHODOLOGY/PRINCIPAL FINDINGS: We used empirical data and inverse modeling--a technique developed by plant ecologists to model seed dispersal--to quantify and compare the dispersal kernels of queens from three Amazonian ant species that compete for access to host-plants. We found that the modal colonization distance of queens varied 8-fold, with the generalist ant species (Crematogaster laevis) having a greater modal distance than two specialists (Pheidole minutula, Azteca sp.) that use the same host-plants. However, our results also suggest that queens of Azteca sp. have maximal distances that are four-sixteen times greater than those of its competitors. CONCLUSIONS/SIGNIFICANCE: We found large differences between ant species in both the modal and maximal distance ant queens disperse to find vacant seedlings used to found new colonies. These differences could result from interspecific differences in queen body size, and hence wing musculature, or because queens differ in their ability to identify potential host plants while in flight. Our results provide support for one of the necessary conditions underlying several of the hypothesized mechanisms promoting coexistence in tropical plant-ants. They also suggest that for some ant species limited dispersal capability could pose a significant barrier to the rescue of populations in isolated forest fragments. Finally, we demonstrate that inverse models parameterized with field data are an excellent means of quantifying the dispersal of ant queens.

Rapid assessment of fruit-color selection by birds using artificial fruits at local scale in Central Amazonia / Utilização de frutos artificiais para avaliar seleção de cores por aves em escala local na Amazônia Central

Modeling clays have been used in several ecological experiments and have proved to be an important tool to variables control. The objective of our study was to determine if fruit color in isolated and grouped displays influences the fruit selection by birds in the field using artificial fruits. Data were collected in six plots distributed homogeneously in 3 km long trails with a minimum distance of 0.5 km. We used a paired experimental design to establish our experiments, so that all treatments were available to the local bird community in each plot. Overall, red was more pecked than brown and white. Isolated red and brown displays were significantly more pecked than others display. Even though our study was conducted in small spatial scales, artificial fruits appeared to be efficient in register fruit consumption attempts by bird. Although inconclusive about selective forces that sharp the dynamics of fruit color polymorphisms and choice by frugivorous birds, our findings corroborate recent studies wherein birds showed preferences by high- over low-contrast fruit signals.

Atualmente o uso de massa de modelar tem sido amplamente utilizado para testar várias hipóteses ecológicas. O objetivo deste trabalho foi determinar se cores de frutos artificiais dispostos isoladamente nos ramos, ou em agregação, influenciaram a escolha das aves. Coletamos os dados em seis parcelas distribuídas homogeneamente em uma trilha de 3 km. Nós usamos desenho experimental pareado, de modo que todos os tratamentos estavam disponíveis para a comunidade local de aves em cada parcela. Os frutos artificiais vermelhos foram mais bicados do que marrons e brancos. Separando-se nos dois diferentes arranjos, vermelhos e marrons isolados foram significativamente mais bicados de que os outros arranjos. Embora nosso estudo tenha sido realizado em pequena escala espacial, frutos artificiais foram eficientes em registrar tentativas de consumo pelas aves. Embora não conclusivo a respeito da pressão seletiva que molda a dinâmica de polimorfismos nas cores dos frutos, bem como dos padrões de escolha pelos frugívoros, nosso trabalho corrobora recentes estudos onde aves preferiram frutos com maior contraste em relação à coloração de fundo imposta pela vegetação.

Interaction intimacy affects structure and coevolutionary dynamics in mutualistic networks.

The structure of mutualistic networks provides clues to processes shaping biodiversity [1-10]. Among them, interaction intimacy, the degree of biological association between partners, leads to differences in specialization patterns [4, 11] and might affect network organization [12]. Here, we investigated potential consequences of interaction intimacy for the structure and coevolution of mutualistic networks. From observed processes of selection on mutualistic interactions, it is expected that symbiotic interactions (high-interaction intimacy) will form species-poor networks characterized by compartmentalization [12, 13], whereas nonsymbiotic interactions (low intimacy) will lead to species-rich, nested networks in which there is a core of generalists and specialists often interact with generalists [3, 5, 7, 12, 14]. We demonstrated an association between interaction intimacy and structure in 19 ant-plant mutualistic networks. Through numerical simulations, we found that network structure of different forms of mutualism affects evolutionary change in distinct ways. Change in one species affects primarily one mutualistic partner in symbiotic interactions but might affect multiple partners in nonsymbiotic interactions. We hypothesize that coevolution in symbiotic interactions is characterized by frequent reciprocal changes between few partners, but coevolution in nonsymbiotic networks might show rare bursts of changes in which many species respond to evolutionary changes in a single species.

Cheating the cheater: domatia loss minimizes the effects of ant castration in an Amazonian ant-plant.

We studied the relationship between Hirtella myrmecophila (Chrysobalanaceae), a common but little-studied Amazonian ant-plant that produces leaf-pouches as domatia, and its obligate ant partner, Allomerus octoarticulatus. Field observations revealed that H. myrmecophila drops domatia from older leaves, a characteristic that is unique among myrmecophytes. The physiological mechanism for abortion of domatia is currently unknown, but this characteristic allows for the existence, within the same plant, of branches with and without ants. Older branches generally bear only old leaves with no domatia and therefore have no ants, whereas younger branches have leaves of various ages. Ants forage mainly on new leaves, and experimental removal of ants showed that A. octoarticulatus is crucial for defense of these leaves against insect herbivores. However, A. octoarticulatus also acts as a castration parasite, severing the plant's inflorescences. Mature flowers and fruits were only found on older branches with no ants, and flower production was 8 times greater on plants whose ants were experimentally removed than on control plants. Given the reproductive costs inflicted by its mutualistic partner, we suggest that abortion of domatia is a strategy developed by H. myrmecophila to minimize the effects of cheating by A. octoarticulatus. These results support the view that evolutionary conflicts of interest between mutualistic species often impose selection for cheating on the partner, as well as for mechanisms to retaliate or to prevent super-exploitation. Opposing selection pressures, operating independently on the two partners, probably help to maintain the evolutionary stability of this mutualistic relationship.