Restricted dispersal reduces the strength of spatial density dependence in a tropical bird population

Spatial processes could play an important role in density-dependent population regulation because the disproportionate use of poor quality habitats as population size increases is widespread in animal populations-the so-called buffer effect. While the buffer effect patterns and their demographic consequences have been described in a number of wild populations, much less is known about how dispersal affects distribution patterns and ultimately density dependence. Here, we investigated the role of dispersal in spatial density dependence using an extraordinarily detailed dataset from a reintroduced Mauritius kestrel (Falco punctatus) population with a territorial (despotic) breeding system. We show that recruitment rates varied significantly between territories, and that territory occupancy was related to its recruitment rate, both of which are consistent with the buffer effect theory. However, we also show that restricted dispersal affects the patterns of territory occupancy with the territories close to release sites being occupied sooner and for longer as the population has grown than the territories further away. As a result of these dispersal patterns, the strength of spatial density dependence is significantly reduced. We conclude that restricted dispersal can modify spatial density dependence in the wild, which has implications for the way population dynamics are likely to be impacted by environmental change.     

Understanding testosterone variation in a tropical lek-breeding bird

Male reproductive coalitions, in which males cooperate to attract females, are a rare strategy among vertebrates. While some studies have investigated ultimate aspects of these relationships, little is known about the mechanistic role that hormones play in modulating cooperative behaviours. Here, we examined male testosterone variation in a tropical lekking bird, the wire-tailed manakin (Pipra filicauda), which exhibits cooperative male-male display coalitions. We found that testosterone levels in territorial males were comparable to those of temperate breeding birds, a surprising result given their environmental, social and reproductive dynamics. In addition, social status rather than plumage was a strong predictor of testosterone variation. Territorial males had significantly higher testosterone levels than did two other plumage classes of floater males, who do not hold territories. We hypothesize that testosterone variation plays an important role in the establishment of male dominance hierarchies (competition), while concurrently facilitating stable display partnerships (cooperation).     

Social and spatial effects on genetic variation between foraging flocks in a wild bird population

Social interactions are rarely random. In some instances, animals exhibit homophily or heterophily, the tendency to interact with similar or dissimilar conspecifics, respectively. Genetic homophily and heterophily influence the evolutionary dynamics of populations, because they potentially affect sexual and social selection. Here, we investigate the link between social interactions and allele frequencies in foraging flocks of great tits (Parus major) over three consecutive years. We constructed co‐occurrence networks which explicitly described the splitting and merging of 85,602 flocks through time (fission–fusion dynamics), at 60 feeding sites. Of the 1,711 birds in those flocks, we genotyped 962 individuals at 4,701 autosomal single nucleotide polymorphisms (SNPs). By combining genomewide genotyping with repeated field observations of the same individuals, we were able to investigate links between social structure and allele frequencies at a much finer scale than was previously possible. We explicitly accounted for potential spatial effects underlying genetic structure at the population level. We modelled social structure and spatial configuration of great tit fission–fusion dynamics with eigenvector maps. Variance partitioning revealed that allele frequencies were strongly affected by group fidelity (explaining 27%–45% of variance) as individuals tended to maintain associations with the same conspecifics. These conspecifics were genetically more dissimilar than expected, shown by genomewide heterophily for pure social (i.e., space‐independent) grouping preferences. Genomewide homophily was linked to spatial configuration, indicating spatial segregation of genotypes. We did not find evidence for homophily or heterophily for putative socially relevant candidate genes or any other SNP markers. Together, these results demonstrate the importance of distinguishing social and spatial processes in determining population structure.     

Breeding bird diversity in relation to environmental gradients in China

Geographic variation in species richness has been explained by different theories such as energy, productivity, energy–water balance, habitat heterogeneity, and freezing tolerance. This study determines which of these theories best account for gradients of breeding bird richness in China. In addition, we develop a best-fit model to account for the relationship between breeding bird richness and environment in China. Breeding bird species richness in 207 localities (3271 km² per locality on average) from across China was related to thirteen environmental variables after accounting for sampling area. The Akaike's information criterion (AIC) was used to evaluate model performance. We used Moran's I to determine the magnitude of spatial autocorrelation in model residuals, and used simultaneous autoregressive model to determine coefficients of determination and AIC of explanatory variables after accounting for residual spatial autocorrelation. Of all environmental variables examined, normalized difference vegetation index, a measure of plant productivity, is the best variable to explain the variance in breeding bird richness. We found that species richness of breeding birds at the scale examined is best predicted by a combination of plant productivity, elevation range, seasonal variation in potential evapotranspiration, and mean annual temperature. These variables explained 47.3% of the variance in breeding bird richness after accounting for sampling area; most of the explained variance in richness is attributable to the first two of the four variables.     

Phenotypic variation and spatial structure of secondary chemistry in a natural population of a tropical tree species

To understand herbivore selection in natural plant populations, it is important to understand the landscape of plant chemical phenotypes that herbivores face and the sources of variation that will define this landscape. We studied the spatial patterns of variation in leaf secondary chemistry of the tropical tree Quararibea asterolepis , Pitt. (Bombacaceae) in a natural population on Barro Colorado Island, Panama, and used this background to discuss hypotheses of natural selection by herbivores. Quararibea plants collected from different sites had consistent differences in their chemical phenotypes. Some of these differences were explained by developmental and environmental sources of variation. Canopy trees had 13% lower yield of leaf extracts than gap seedlings, explained by 41% lower concentrations of the more abundant metabolites in the secondary compound profile. Also, plants growing in gaps had 25% higher yield than those in the understory, explained by two-fold increases in the concentration of some of the less polar secondary compounds in the profile. Differences in soil type did not affect the secondary chemistry of leaves, but sites with different topography had differences in the secondary compound profile that were not explained by any of the measured environmental sources of variation. Neighboring parent-offspring pairs and sibling/half sibling clusters displayed equal or higher variance among themselves than unrelated individuals at farther distances. Assuming that related plants should be more similar in their phenotypes, this pattern is consistent with local selection by herbivores overriding the similarity of related plants in a frequency- or distance-dependent manner.     

Assessing the influence of environmental gradients on seed mass variation in mountain grasslands using a spatial phylogenetic filtering approach

Several studies have demonstrated that seed mass is related to different environmental factors. However, they have taken no account of the joint effects of spatial and phylogenetic information. We analysed the distribution pattern of seed mass along an elevational gradient (1040–2380 m a.s.l.) at the community level in grasslands of the southern Alps. First, we tested the influence of environmental filters (climate and soil properties) in determining community-weighted seed mass variation in mountain grasslands. Second, we verified the relative roles of environmental filters in determining seed mass variation after accounting for spatial and phylogenetic autocorrelation with an eigenvector filtering approach. Temperature, soil fertility, and soil pH were the most important predictors for explaining seed mass variation; specifically, warmer, low fertility, and alkaline grasslands showed a greater seed mass. Inclusion of spatio-phylogenetic filters in the model increased its fit and the variance explained and reduced autocorrelation significantly but had substantial effects on the parameter estimates, with temperature and soil pH becoming insignificant. This effect may be ascribable to spatially structured phylogenetic patterns and could likely result from the common evolutionary histories shared by many species at sites with similar environmental conditions. Therefore, the observed patterns between community-weighted seed mass and both temperature and soil pH are not independent of phylogeny, but they are explained by the shared history within genera and families. Nevertheless, soil fertility remained the most important predictor for explaining seed mass variation. The results of this work contribute to better understanding the combined effects of environment and evolutionary factors for determining seed mass distributions in the spatial context of mountain grasslands. The observed relationships with climate and soil properties are particularly interesting because they are potentially relevant when modelling plant trait composition under changes in land use and climate.     

The genetic basis of spatial cognitive variation in a food-caching bird

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Disentangling the spatial and temporal causes of decline in a bird population

The difficulties in understanding the underlying reasons of a population decline lie in the typical short duration of field studies, the often too small size already reached by a declining population or the multitude of environmental factors that may influence population trend. In this difficult context, useful demographic tools such as integrated population models (IPM) may help disentangling the main reasons for a population decline. To understand why a hoopoe Upupa epops population has declined, we followed a three step model analysis. We built an IPM structured with respect to habitat quality (approximated by the expected availability of mole crickets, the main prey in our population) and estimated the contributions of habitat‐specific demographic rates to population variation and decline. We quantified how much each demographic rate has decreased and investigated whether habitat quality influenced this decline. We tested how much weather conditions and research activities contributed to the decrease in the different demographic rates. The decline of the hoopoe population was mainly explained by a decrease in first‐year apparent survival and a reduced number of fledglings produced, particularly in habitats of high quality. Since a majority of pairs bred in habitats of the highest quality, the decrease in the production of locally recruited yearlings in high‐quality habitat was the main driver of the population decline despite a homogeneous drop of recruitment across habitats. Overall, the explanatory variables we tested only accounted for 19% of the decrease in the population growth rate. Among these variables, the effects of spring temperature (49% of the explained variance) contributed more to population decline than spring precipitation (36%) and research activities (maternal capture delay, 15%). This study shows the power of IPMs for identifying the vital rates involved in population declines and thus paves the way for targeted conservation and management actions.     

Seasonal variation in primary productivity in three tropical ponds

Summary Primary productivity values ranged from 1.5 to 15.8 gc/M²/day in Othakadai pond; 2.0 to 8.0 gc/M²/day in Teppakulam tank and 1.05 to 5.4 gc/M²/day in Yanamalai pond. Among the three ponds, highest annual yield was recorded in Othakadai pond (2000.2 gc/M²/yr) next in order comes Teppakulam tank (1211.8 gc/M²/yr) and lastly Yanamalai pond (810.7 gc/M²/yr). In all three ponds temperature and chlorophyll-a showed a more or less direct correlation to productivity values. Alkalinity values and productivity values showed a positive correlation in Othakadai and Yanamalai ponds, while no relationship between alkalinity and productivity could be established in Teppakulam tank.Formed part of a Ph. D. Thesis submitted to Madurai University.     

Explaining variation in tropical plant community composition: influence of environmental and spatial data quality

The degree to which variation in plant community composition (beta-diversity) is predictable from environmental variation, relative to other spatial processes, is of considerable current interest. We addressed this question in Costa Rican rain forest pteridophytes (1,045 plots, 127 species). We also tested the effect of data quality on the results, which has largely been overlooked in earlier studies. To do so, we compared two alternative spatial models [polynomial vs. principal coordinates of neighbour matrices (PCNM)] and ten alternative environmental models (all available environmental variables vs. four subsets, and including their polynomials vs. not). Of the environmental data types, soil chemistry contributed most to explaining pteridophyte community variation, followed in decreasing order of contribution by topography, soil type and forest structure. Environmentally explained variation increased moderately when polynomials of the environmental variables were included. Spatially explained variation increased substantially when the multi-scale PCNM spatial model was used instead of the traditional, broad-scale polynomial spatial model. The best model combination (PCNM spatial model and full environmental model including polynomials) explained 32% of pteridophyte community variation, after correcting for the number of sampling sites and explanatory variables. Overall evidence for environmental control of beta-diversity was strong, and the main floristic gradients detected were correlated with environmental variation at all scales encompassed by the study (c. 100–2,000 m). Depending on model choice, however, total explained variation differed more than fourfold, and the apparent relative importance of space and environment could be reversed. Therefore, we advocate a broader recognition of the impacts that data quality has on analysis results. A general understanding of the relative contributions of spatial and environmental processes to species distributions and beta-diversity requires that methodological artefacts are separated from real ecological differences.     

Nonrandom variation of morphological traits across environmental gradients in a land snail

Morphological variation is often attributed to differential adaptations to diverse habitats, but adaptations to a similar environment do not necessarily result in similar phenotypes. Adaptations for water and heat budget are crucial for organisms living in arid habitats, and in snails, variation in shell morphology has been frequently attributed to selection by stressful environmental factors. However, their phenotypic divergence often is not accompanied by a relevant niche differentiation and consistent relationships with environmental correlates are lacking. In the pulmonate genus Albinaria, there is great size and shape variation between and within species, and there are two major shell sculpture morphotypes, ribbed and smooth. We used 62 populations of 28 Albinaria species, taking into account their phylogeny, to examine the variation of shell traits (sculpture, size, shape), their effect on water and heat budget, and their association with geographical and climatic gradients. We found unambiguous size and shape discrimination between the two morphotypes. Ribbed shells are lighter, taller, and slimmer and have a smaller aperture than the smooth ones. Moreover, significant correlations between shell traits and heat/moisture budget and climate/geography were revealed. Ribbed and taller shells retain more water on their shell surface, and on the other hand, smooth shells exhibit lower water permeability. Therefore, two strategies are being used to prevent water loss, active retention or resistance to loss. Consequently, different alternative solutions evolved and were retained as responses to the same stressful factor by the two distinct shell morphotypes. Larger shells occur in southern latitudes, mostly on islands, and at sites where there is a shortage of rainfall. Therefore, the variation of the examined traits is nonrandom with respect to location and to climate and their evolution can be attributed to selection by environmental factors, with water availability being the key driving agent of body-size variation.     

Phylogenetic turnover along local environmental gradients in tropical forest communities

Oecologia - While the importance of local-scale habitat niches in shaping tree species turnover along environmental gradients in tropical forests is well appreciated, relatively little is known...     

Multiple density-dependence mechanisms regulate a migratory bird population during the breeding season

The mechanisms regulating bird populations are poorly understood and controversial. We provide evidence that a migratory songbird, the black-throated blue warbler (Dendroica caerulescens), is regulated by multiple density-dependence mechanisms in its breeding quarters. Evidence of regulation includes: stability in population density during 1969-2002, strong density dependence in time-series analyses of this period, an inverse relationship between warbler density and annual fecundity, and a positive relationship between annual fecundity and recruitment of yearlings in the subsequent breeding season. Tests of the mechanisms causing regulation were carried out within the Hubbard Brook Experimental Forest, New Hampshire, during 1997-1999. When individuals from abutting territories were experimentally removed in a homogeneous patch of high-quality habitat, the fecundity of focal pairs nearly doubled, revealing a locally operating crowding mechanism. A site-dependence mechanism was indicated by an inverse relationship between population size and mean territory quality, as well as by greater annual fecundity on the sites that were most frequently occupied and of highest quality. These site-dependence relationships were revealed by intensive monitoring of territory quality and demography at the landscape spatial scale. Crowding and site-dependence mechanisms, therefore, acted simultaneously but at different spatial scales to regulate local abundance of this migratory bird population.     

Dispersal and spatial scale affect synchrony in spatial population dynamics

A large body of theoretical studies has shown that synchrony among populations is critical for the long-term persistence of species in fragmented habitats. Although the effects of dispersal and environmental factors on synchrony have been investigated theoretically, empirical studies of these relationships have been lacking. We explored the interplay between environmental and demographic factors (fecundity, survival, dispersal) on population synchrony for 53 species of birds. We show that the interspecific differences in mean synchrony were determined by global environmental factors whose action was probably mediated by the abundance of each species. After removing the effect of these global factors on synchrony, the residual synchrony was strongly correlated with dispersal distance. The relationship between dispersal and synchrony was stronger for the species nesting in wet habitats than for those nesting in dry habitats. Our results indicate that different factors synchronize bird populations at different spatial scales, thus highlighting the role of scale in understanding spatial population dynamics and extinction risks.     

Quantifying how fine-grained environmental heterogeneity and genetic variation affect demography in an annual plant population

The ability of plant species to colonize new habitats and persist in changing environments depends on their ability to respond plastically to environmental variation and on the presence of genetic variation, thus allowing adaptation to new conditions. For invasive species in particular, the relationship between phenotypic trait expression, demography, and the quantitative genetic variation that is available to respond to selection are likely to be important determinants of the successful establishment and persistence of populations. However, the magnitude and sources of individual demographic variation in exotic plant populations remain poorly understood. How important is plasticity versus adaptability in populations of invasive species? Among environmental factors, is temperature, soil nutrients, or competition most influential, and at what scales and life stages do they affect the plants? To investigate these questions we planted seeds of the exotic annual plant Erodium brachycarpum into typical pasture habitat in a spatially nested design. Seeds were drawn from 30 inbred lines to enable quantification of genetic effects. Despite a positive population growth rate, a few plants (0.1?%) produced >50?% of the seeds, suggesting a low effective population size. Emergence and early growth varied by genotype, but as in previous studies on native plants, environmental effects greatly exceeded genetic effects, and survival was unrelated to genotype. Environmental influences shifted from microscale soil compaction and litter depth at emergence through to larger-scale soil nutrient gradients during growth and to competition during later survival and seed production. Temperature had no effect. Most demographic rates were positively correlated, but emergence was negatively correlated with other rates.     

Habitat fragmentation differentially shapes neutral and immune gene variation in a tropical bird species

Habitat fragmentation is a major cause of biodiversity loss, responsible for an alteration of intraspecific patterns of neutral genetic diversity and structure. Although neutral genetic variation can be informative for demographic inferences, it may be a poor predictor of adaptive genetic diversity and thus of the consequences of habitat fragmentation on selective evolutionary processes. In this context, we contrasted patterns of genetic diversity and structure of neutral loci (microsatellites) and immune genes (i.e., toll-like receptors) in an understorey bird species, the wedge-billed woodcreeper Glyphorynchus spirurus. The objectives were (1) to investigate forest fragmentation effects on population genetic diversity, (2) to disentangle the relative role of demography (genetic drift and migration) and selection, and (3) to assess whether immunogenetic patterns could be associated with variation of ectoparasite (i.e., ticks) pressures. Our results revealed an erosion of neutral genetic diversity and a substantial genetic differentiation among fragmented populations, resulting from a decrease in landscape connectivity and leading to the divergence of distinct genetic pools at a small spatial scale. Patterns of genetic diversity observed for TLR4 and TLR5 were concordant with neutral genetic patterns, whereas those observed for TLR3 and TLR21 were discordant. This result underlines that the dominant evolutionary force shaping immunogenetic diversity (genetic drift vs. selection) may be different depending on loci considered. Finally, tick prevalence was higher in fragmented environments. We discussed the hypothesis that pathogen selective pressures may contribute to maintain adaptive genetic diversity despite the negative demographic effect of habitat fragmentation on neutral genetic diversity.Subject terms: Tropical ecology, Genetic variation     

Climate change and the risks associated with delayed breeding in a tropical wild bird population

There is growing evidence of changes in the timing of important ecological events, such as flowering in plants and reproduction in animals, in response to climate change, with implications for population decline and biodiversity loss. Recent work has shown that the timing of breeding in wild birds is changing in response to climate change partly because individuals are remarkably flexible in their timing of breeding. Despite this work, our understanding of these processes in wild populations remains very limited and biased towards species from temperate regions. Here, we report the response to changing climate in a tropical wild bird population using a long-term dataset on a formerly critically endangered island endemic, the Mauritius kestrel. We show that the frequency of spring rainfall affects the timing of breeding, with birds breeding later in wetter springs. Delays in breeding have consequences in terms of reduced reproductive success as birds get exposed to risks associated with adverse climatic conditions later on in the breeding season, which reduce nesting success. These results, combined with the fact that frequency of spring rainfall has increased by about 60 per cent in our study area since 1962, imply that climate change is exposing birds to the stochastic risks of late reproduction by causing them to start breeding relatively late in the season.     

Top-down response to spatial variation in productivity and bottom-up response to temporal variation in productivity in a long-term study of desert ants

Under the Ecosystem Exploitation Hypothesis ecosystem productivity predicts trophic complexity, but it is unclear if spatial and temporal drivers of productivity have similar impacts. Long-term studies are necessary to capture temporal impacts on trophic structure in variable ecosystems such as deserts. We sampled ants and measured plant resources in the Simpson Desert, central Australia over a 22-year period, during which rainfall varied 10-fold. We sampled dune swales (higher nutrient) and crests (lower nutrient) to account for spatial variation in productivity. We asked how temporal and spatial variation in productivity affects the abundance of ant trophic guilds. Precipitation increased vegetation cover, with the difference more pronounced on dune crests; seeding and flowering also increased with precipitation. Generalist activity increased over time, irrespective of productivity. Predators were more active in more productive (swale) habitat, i.e. spatial impacts of productivity were greatest at the highest trophic level. By contrast, herbivores (seed harvesters and sugar feeders) increased with long-term rainfall; seed harvesters also increased as seeding increased. Temporal impacts of productivity were therefore greatest for low trophic levels. Whether productivity variation leads to top-down or bottom-up structured ecosystems thus depends on the scale and dimension (spatial or temporal) of productivity.