Intra‐ and inter‐individual variation show distinct trends as drivers of seasonal changes in the resource use of a neotropical marsupial

Individual specialization can influence important ecological and evolutionary traits and both inter‐ and intra‐individual variation in resource use can drive niche shifts in natural populations. We evaluated the predominance of these two factors for determining seasonal differences in the trophic niche of the didelphid marsupial Gracilinanus agilis (Burmeister, 1854) in the highly seasonal Brazilian savanna. In the three sampled sites, the population of G. agilis increased its dietary niche width in the warm–wet season, when food resources are more abundant, and there were no differences between sexes and no interaction between season and sex. However, the evaluation of intra‐individual variation indicated that females reduce the number of items consumed during the warm–wet season, whereas males show no seasonal differences. Inter‐individual variation nonetheless followed the overall population pattern because both sexes increased their spread with respect to food‐item consumption in the warm–wet season. Additionally, we found positive relationships between body length and diet only in the warm–wet season, when larger animals fed more on invertebrates and less on fruits than the small ones. Our results show a previously unknown pattern for mammals, in which the trophic niche is wider during the high‐resource season as a result of inter‐individual variation along the body‐size axis. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 737–747.     

Seasonal population and individual niche dynamics in a tetra fish in the Pantanal wetlands

In seasonal tropical regions, rainfall and/or temporary floods during the wet season generally increase the abundance and diversity of food resources to many consumers as compared to the dry season. Therefore, seasonality can affect intraspecific competition and ecological opportunity, which are two important ecological mechanisms underlying population and individual niche variations. Here, we took advantage of the strong seasonality in the Pantanal wetlands to investigate how within‐ and between‐individual diet variations relate to seasonal population niche dynamics of the tetra fish Astyanax lacustris. We quantified dietary niche using gut contents and stable isotopes. Tetras had higher gut fullness and better body condition in the wet season, suggesting that competition is more intense in the dry season. The population niche was broader in the wet season due to an increase in diet divergence between individuals, in spite of potential stronger competition in the dry season. We posit that low ecological opportunity in the dry season limits the diversifying effect of intraspecific competition, constraining population niche expansion. Our results add new insights on how seasonality affects population and individual diets, indicating that intraspecific competition and ecological opportunity interact to determine temporal niche variations in seasonal environments.     

Individual–resource networks reveal distinct fruit preferences of selective individuals from a generalist population of the Helmeted Manakin

Ecological studies traditionally assume that generalist populations are homogeneous in the use of food resources, but empirical evidence supports that intraspecific differences in morphology, physiology and behaviour affect foraging decisions and promote diet variation among individuals. Furthermore, the temporal availability of resources may shape the dynamics of population trophic niche, which ultimately depends on individual niches. In this study, we investigated the seasonal changes in individual-based networks between the Helmeted Manakin Antilophia galeata, a generalist frugivorous bird, and fruiting plants, following theoretical models of interindividual diet variation based on the Optimal Diet Theory. Selective individuals were the majority of the generalist population of the Helmeted Manakin. Our results suggest that the structure of the individual-resource networks varied seasonally. We found that modularity was higher than expected by chance in the wet season, when fruit availability was also higher. In the dry season, modules disappeared and the network became more nested. These findings are consistent with the Distinct Preference Model of diet variation. We suggest that downscaling ecological networks to the individual level may reveal emergent properties that, albeit existent, are not evident in species–resources networks.     

Availability of food resources and habitat structure shape the individual‐resource network of a Neotropical marsupial

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Trophic niche, age structure and seasonality in Dolichopoda cave crickets

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Seasonal differences in population‐, ensemble‐ and community‐level responses of bats to landscape structure in Amazonia

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Individual variation in resource use by opossums leading to nested fruit consumption

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Resource diversity promotes among‐individual diet variation,but not genomic diversity,in lake stickleback

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Beyond ecological opportunity: Prey diversity rather than abundance shapes predator niche variation

1. Ecological opportunity (i.e. the diversity of available resources) has a pivotal role in shaping niche variation and trophic specialisation of animals. However, ecological opportunity can be described with regard to both diversity and abundance of resources. The degree to which these two components contribute to niche variation remains unexplored.
2. To address this, we used an extensive dataset on fish diet and benthic invertebrate diversity and density from 73 sampling events in three Norwegian rivers in order to explore realised trophic niches and the response of dietary niche variation along gradients of resource diversity (potential trophic niches), resource density (as a proxy of resource abundance) and fish density (as a proxy of inter‐ and intra‐specific competition) in a freshwater top predator (the brown trout, Salmo trutta L.).
3. Linear models indicated that individual and population niche variation increased with increasing ecological opportunity in terms of prey diversity. However, no simple cause‐and‐effect associations between niche indices and prey abundance were found. Our multiple regression analyses indicated that the abundance of certain resources (e.g. Chironomidae) can interact with prey diversity to determine individual and population realised trophic niches. Niche variation (within‐individual component and inter‐individual diet variation) decreased with increasing inter‐ and intra‐specific competition.
4. This study extends prevailing trophic ecology theory by identifying diversity, rather than density, of available prey resources as a primary driver of niche variation in fish of temperate riverine systems with no extensive resource limitation. The study also shows that ecological opportunity may mask the direction of the effect (compression or expansion) of competition on niche variation when food resources are diverse.
5. Our study provides novel empirical insight to the driving forces behind niche variation and reveals that diversity, rather than density, of available prey resources may be a primary driver of niche variation in freshwater fish. Our study supports the view that a broader potential trophic niche promotes broader realised trophic niche variation by individuals, which leads to individual niche diversification by opening access to alternatives resources, resulting in a concomitant rise in the realised trophic niche width of the population.

     

The seasonal dynamic of ant‐flower networks in a semi‐arid tropical environment

1. Several studies have recently focused on the structure of ecological networks involving ants and plants with extrafloral nectaries; however, little is known about the effects of temporal variation in resource abundance on the structure of ant–plant networks mediated by floral nectar. 2. In this study, it was evaluated how strong seasonality in resource availability in a semi‐arid tropical environment affects the structure of ant–flower networks. We recorded ants collecting floral nectar during two seasons (from December 2009 to January 2013): dry and green seasons. Then, we built interaction networks for flower‐visiting ants in the Brazilian Caatinga separately for each combination of transect and season. 3. In general, strong seasonality directly influenced patterns of ant–flower interactions and the overall complexity of these ecological networks. During the dry season, networks were more connected, less modular, and exhibited greater niche overlap of flower‐visiting ants than during the green season. Moreover, resource utilisation by ants during the dry season tended to be more aggregated. These findings indicate that during the dry season, ant species tended to share many resource bases, probably owing to lower overall resource availability during this season. Species composition of the ant network component was highly season specific; however, a central core of highly generalised ants was present during both seasons. 4. The stability of this central core between seasons could strongly affect the ecological and evolutionary dynamics of these interaction networks. This study contributes to the understanding of the structure and dynamics of ant‐flower interactions in extremely seasonal environments.     

Season‐modulated responses of Neotropical bats to forest fragmentation

Seasonality causes fluctuations in resource availability, affecting the presence and abundance of animal species. The impacts of these oscillations on wildlife populations can be exacerbated by habitat fragmentation. We assessed differences in bat species abundance between the wet and dry season in a fragmented landscape in the Central Amazon characterized by primary forest fragments embedded in a secondary forest matrix. We also evaluated whether the relative importance of local vegetation structure versus landscape characteristics (composition and configuration) in shaping bat abundance patterns varied between seasons. Our working hypotheses were that abundance responses are species as well as season specific, and that in the wet season, local vegetation structure is a stronger determinant of bat abundance than landscape‐scale attributes. Generalized linear mixed‐effects models in combination with hierarchical partitioning revealed that relationships between species abundances and local vegetation structure and landscape characteristics were both season specific and scale dependent. Overall, landscape characteristics were more important than local vegetation characteristics, suggesting that landscape structure is likely to play an even more important role in landscapes with higher fragment‐matrix contrast. Responses varied between frugivores and animalivores. In the dry season, frugivores responded more to compositional metrics, whereas during the wet season, local and configurational metrics were more important. Animalivores showed similar patterns in both seasons, responding to the same group of metrics in both seasons. Differences in responses likely reflect seasonal differences in the phenology of flowering and fruiting between primary and secondary forests, which affected the foraging behavior and habitat use of bats. Management actions should encompass multiscale approaches to account for the idiosyncratic responses of species to seasonal variation in resource abundance and consequently to local and landscape scale attributes.     

Dietary overlap and seasonality in three species of mormoopid bats from a tropical dry forest

Competing hypotheses explaining species’ use of resources have been advanced. Resource limitations in habitat and/or food are factors that affect assemblages of species. These limitations could drive the evolution of morphological and/or behavioural specialization, permitting the coexistence of closely related species through resource partitioning and niche differentiation. Alternatively, when resources are unlimited, fluctuations in resources availability will cause concomitant shifts in resource use regardless of species identity. Here, we used next‐generation sequencing to test these hypotheses and characterize the diversity, overlap and seasonal variation in the diet of three species of insectivorous bats of the genus Pteronotus. We identified 465 prey (MOTUs) in the guano of 192 individuals. Lepidoptera and Diptera represented the most consumed insect orders. Diet of bats exhibited a moderate level of overlap, with the highest value between Pteronotus parnellii and Pteronotus personatus in the wet season. We found higher dietary overlap between species during the same seasons than within any single species across seasons. This suggests that diets of the three species are driven more by prey availability than by any particular predator‐specific characteristic. P. davyi and P. personatus increased their dietary breadth during the dry season, whereas P. parnellii diet was broader and had the highest effective number of prey species in all seasons. This supports the existence of dietary flexibility in generalist bats and dietary niche overlapping among groups of closely related species in highly seasonal ecosystems. Moreover, the abundance and availability of insect prey may drive the diet of insectivores.     

The role of El Niño–Southern Oscillation in the dynamics of a savanna large herbivore population

Our understanding of large‐scale climatic phenomena and dynamics of large herbivore populations comes principally from research in northern regions with temperate, seasonal climate and animal communities with relatively low species diversity. To assess the generality of that perspective, we investigated effects of El Niño–Southern Oscillation (ENSO) on population dynamics of African buffalo Syncerus caffer inhabiting a semi‐arid savanna with variable rainfall. We used linear and nonlinear‐threshold models to investigate relationships between population parameters and explanatory variables affecting forage conditions (seasonal rainfall, Southern Oscillation Index [SOI]). El Niño‐related droughts in 1982–1983 and 1991–1992 were associated with strongly negative population change, a pattern expected to coincide with a decrease in normally high and constant adult survival. Consistent with that nonlinear pattern, we detected threshold relationships between wet‐season rainfall and population change. Juvenile recruitment was described best by linear relationships with dry‐season. Because ENSO operates primarily through wet‐season rainfall, whereas population dynamics were also related to dry‐season rainfall, SOI did not have the predictive ability of individual weather components.     

Dietary niche and population dynamic feedbacks in a novel habitat

Population dynamics and resource use are often intricately connected via density‐dependent intraspecific competition. However, experimental studies of concurrent change in population and resource use dynamics are scarce. In particular, the impact of factors such as genetic diversity, which can affect both population dynamics and competition, remains unexplored. Using stable isotope analysis and periodic population censuses, we quantified both diet and population dynamics in wheat‐adapted Tribolium castaneum (flour beetle) populations provided with an additional novel resource (corn). Populations were initiated with different levels of genetic variation for traits relevant to population growth and resource use (e.g. fecundity and survival).We found that high population size decreased subsequent corn use, and high corn use in turn lowered population size. Surprisingly, we did not detect a significant effect of founding genetic variation on resource niche expansion, although genetic variation increased overall population size and stability. In contrast, dietary niche expansion decreased both population size and stability. Finally, larval and adult niche dynamics were uncorrelated, suggesting that various life stages perceive or respond differentially to intraspecific competition and resource availability. Our experiments indicate that population performance in a novel habitat depends on stage‐specific interactions between resource use, standing genetic variation, and population size.     

Testing the niche variation hypothesis with a measure of body condition

Individual variation and fitness are cornerstones of evolution by natural selection. The niche variation hypothesis (NVH) posits that when interspecific competition is relaxed, intraspecific competition should drive niche expansion by selection favoring use of novel resources and that among‐individual variation should confer a selective advantage. Population‐level niche expansion could be achieved by all individuals using all available resources, or by each individual using a unique combination of resources, thereby increasing among‐individual dietary niche variation. Although individual variation can lead to species‐level evolutionary and ecological change, observed variation does not ensure a beneficial outcome. We used carbon and nitrogen stable isotope analysis of claw keratin and a Bayesian stable isotope mixing model to estimate the summer (July–September) assimilated diet of individual female black Ursus americanus and brown U. arctos bears. We quantified variation in dietary niche in both populations, and assessed diet relative to percentage body fat. We hypothesized that if the NVH held, percentage body fat would be similar for individuals of the same species across much of the dietary range of observed proportional salmon contributions to individual bear diets. Although we found greater differences in dietary niches between than within species, we observed greater among‐individual dietary variation in the brown bear population. Moreover, we found that within each species individual female bears achieved similar ranges of percentage body fat at various levels of salmon in the diet. Our results provide support for the NVH. Linking individual dietary niches to measures of physiological condition related to fitness can offer new insights into eco‐evolutionary processes related to food resource use.     

Population and evolutionary dynamics in spatially structured seasonally varying environments

Increasingly imperative objectives in ecology are to understand and forecast population dynamic and evolutionary responses to seasonal environmental variation and change. Such population and evolutionary dynamics result from immediate and lagged responses of all key life‐history traits, and resulting demographic rates that affect population growth rate, to seasonal environmental conditions and population density. However, existing population dynamic and eco‐evolutionary theory and models have not yet fully encompassed within‐individual and among‐individual variation, covariation, structure and heterogeneity, and ongoing evolution, in a critical life‐history trait that allows individuals to respond to seasonal environmental conditions: seasonal migration. Meanwhile, empirical studies aided by new animal‐tracking technologies are increasingly demonstrating substantial within‐population variation in the occurrence and form of migration versus year‐round residence, generating diverse forms of ‘partial migration’ spanning diverse species, habitats and spatial scales. Such partially migratory systems form a continuum between the extreme scenarios of full migration and full year‐round residence, and are commonplace in nature. Here, we first review basic scenarios of partial migration and associated models designed to identify conditions that facilitate the maintenance of migratory polymorphism. We highlight that such models have been fundamental to the development of partial migration theory, but are spatially and demographically simplistic compared to the rich bodies of population dynamic theory and models that consider spatially structured populations with dispersal but no migration, or consider populations experiencing strong seasonality and full obligate migration. Second, to provide an overarching conceptual framework for spatio‐temporal population dynamics, we define a ‘partially migratory meta‐population’ system as a spatially structured set of locations that can be occupied by different sets of resident and migrant individuals in different seasons, and where locations that can support reproduction can also be linked by dispersal. We outline key forms of within‐individual and among‐individual variation and structure in migration that could arise within such systems and interact with variation in individual survival, reproduction and dispersal to create complex population dynamics and evolutionary responses across locations, seasons, years and generations. Third, we review approaches by which population dynamic and eco‐evolutionary models could be developed to test hypotheses regarding the dynamics and persistence of partially migratory meta‐populations given diverse forms of seasonal environmental variation and change, and to forecast system‐specific dynamics. To demonstrate one such approach, we use an evolutionary individual‐based model to illustrate that multiple forms of partial migration can readily co‐exist in a simple spatially structured landscape. Finally, we summarise recent empirical studies that demonstrate key components of demographic structure in partial migration, and demonstrate diverse associations with reproduction and survival. We thereby identify key theoretical and empirical knowledge gaps that remain, and consider multiple complementary approaches by which these gaps can be filled in order to elucidate population dynamic and eco‐evolutionary responses to spatio‐temporal seasonal environmental variation and change.     

Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist

Longitudinal studies have revealed how variation in resource use within consumer populations can impact their dynamics and functional significance in communities. Here, we investigate multi-decadal diet variations within individuals of a keystone megaherbivore species, the African elephant (Loxodonta africana), using serial stable isotope analysis of tusks from the Kruger National Park, South Africa. These records, representing the longest continuous diet histories documented for any extant species, reveal extensive seasonal and annual variations in isotopic--and hence dietary--niches of individuals, but little variation between them. Lack of niche distinction across individuals contrasts several recent studies, which found relatively high levels of individual niche specialization in various taxa. Our result is consistent with theory that individual mammal herbivores are nutritionally constrained to maintain broad diet niches. Individual diet specialization would also be a costly strategy for large-bodied taxa foraging over wide areas in spatio-temporally heterogeneous environments. High levels of within-individual diet variability occurred within and across seasons, and persisted despite an overall increase in inferred C(4) grass consumption through the twentieth century. We suggest that switching between C(3) browsing and C(4) grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.     

Survival estimation of a cryptic antelope via photographic capture–recapture

Adult survival is a primary determinant of abundance and dynamics of large herbivore populations. For species that are inconspicuous, however, accurate survival estimation depends on accommodating low detection probability. For species with individually recognizable markings, photographic capture–recapture (CR) provides an approach to estimate population parameters while accounting for imperfect detection. I investigated the use of photographic CR for a cryptic large herbivore, the nyala, in a region of Hluhluwe–iMfolozi Park, South Africa. I conducted photographic sampling based on the closed robust design, with 5–6 daily sampling occasions nested within three week‐long sampling periods, which delineated one dry and one wet season. Detection differed between sexes: encounter probability of female adults depended on whether individuals fell into high‐encounter (seasonal range: 0.61–0.71) or low‐encounter (seasonal range: 0.29–0.40) groups, whereas male adults had a constant encounter probability of 0.39 per day. For both sexes, monthly survival probability was ≥0.93 and did not differ appreciably between seasons or sexes. Given the role of survival in population dynamics, photographic CR has the potential to provide survival estimates for cryptic large herbivores that lack such information.     

Seasonal variation in diet breadth of folivorous Lepidoptera in the Brazilian cerrado

Documented patterns of specialization and species interactions often omit plasticity in resource use across space and time, yet such variation is an important part of species interactions. To examine temporal variation in resource use, we compared species‐ and community‐level patterns of host plant use by folivorous caterpillars between the dry and rainy seasons in four preserved areas of cerrado vegetation in the Distrito Federal, Brazil. We sampled plants and caterpillars in 10 m circular plots monthly from March 2010 to March 2011. At the community level, we found a significant increase in the mean diet breadth of dry season caterpillars relative to those collected in the rainy season. Families and species of moths varied in diet breadth, but most exhibited a seasonal expansion. In particular, intraspecific comparisons showed a 30% increase in the number of host plants used in the dry season compared to the rainy season. These results provide a clear example of how temporal variation in resource use is heterogeneous, and more generalized patterns of resource use can emerge from studies at large temporal scales. Thus, seasonal or annual heterogeneity may obscure ecologically relevant specialized interactions that occur at smaller scales.