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Social trematode parasites increase standing army size in areas of greater invasion threat.
Organisms or societies are resource limited, causing important trade-offs between reproduction and defence. Given such trade-offs, optimal allocation theory predicts that, for animal societies with a soldier caste, allocation to soldiers should reflect local external threats. Although both threat intensity and soldier allocation can vary widely in nature, we currently lack strong evidence that spatial variation in threat can drive the corresponding variation in soldier allocation. The diverse guild of trematode parasites of the California horn snail provides a useful system to address this problem. Several of these species form colonies in their hosts with a reproductive division of labour including a soldier caste. Soldiers are non-reproductive and specialized in defence, attacking and killing invading parasites. We quantified invasion threat and soldier allocation for 168 trematode colonies belonging to six species at 26 sites spread among 10 estuaries in temperate and tropical regions. Spatial variation in invasion threat was matched as predicted by the relative number of soldiers for multiple parasite species. Soldier allocation correlated with invasion threat at fine spatial scales, suggesting that allocation is at least partly inducible. These results may represent the first clear documentation of a spatial correlation between allocation to any type of caste and a biotic selective agent
Aquatic beetles influence colonization of disparate taxa in small lentic systems
© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd Structure of natural communities is shaped by both abiotic characteristics and the ongoing processes of community assembly. Important to this process are the habitat selection behaviors and subsequent survival of colonists, both in the context of temporal changes in the abiotic characteristics and priority effects driven by earlier colonists. Aquatic beetles are prevalent in temporary freshwater systems, form speciose assemblages, and are often early colonists of temporary ponds. While beetles have the potential to influence community structure through post-colonization interactions (predation and competition), our goal was to determine whether the presence of beetle assemblages (versus patches without beetles) influences the colonization and oviposition of a diverse group of animals in a naturally colonized experimental landscape. We established mesocosms that either contained existing beetle assemblages or contained no beetles and assessed abundances of subsequent colonists. Treefrogs, Hyla chrysoscelis, and mosquitoes, Culex restuans, both deposited fewer eggs in patches containing beetle assemblages, while two beetles, Copelatus glyphicus and Paracymus, colonized those patches at lower rates. One beetle, Helophorus linearis, colonized patches containing beetle assemblages at higher rates, while two beetles, Berosus infuscatus and Tropisternus lateralis, exhibited no colonization differences between treatments. Overall, there were no differences in the assemblage structure or richness of beetles that colonized patches. Our results illustrate the importance of species-specific habitat selection behavior in determining the species composition of habitat patches, while emphasizing the role of priority effects in influencing patterns of community assembly. Habitat selection in response to abiotic and biotic characteristics of habitat patches can potentially create greater spatiotemporal niche separation among the numerous, often closely related species (phylogenetically and trophically), that can be simultaneously found in similar patches across landscapes
Spatial and Temporal Dynamics of Habitat Selection Across Canopy Gradients Generates Patterns of Species Richness and Composition in Aquatic Beetles
1. Colonisation is a critical ecological process influencing both population and community level dynamics by connecting spatially discrete habitat patches. How communities respond to both natural and anthropogenic disturbances, furthermore, requires a basic understanding of how any environmental change modifies colonisation rates. For example, disturbance-induced shifts in the quantity of forest cover surrounding aquatic habitats have been associated with the distribution and abundance of numerous aquatic taxa. However, the mechanisms generating these broad and repeatable field patterns are unclear. 2. Such patterns of diversity could result from differential spatial mortality post colonisation, or from colonisation alone if species select sites non-randomly along canopy coverage gradients. We examined the colonisation/oviposition dynamics of aquatic beetles in experimental ponds placed under both open and closed forest canopies. 3. Canopy coverage imposed a substantial behavioural filter on the colonisation and reproduction of aquatic beetles representing multiple trophic levels, and resulted in significantly higher abundance, richness, and oviposition activity in open canopy ponds. These patterns strengthened overtime; although early in the experiment, the most abundant beetle had similar abundance in open and closed ponds. However, its abundance subsequently declined and then most other species heavily colonised open canopy ponds. 4. The primary response of many aquatic species to disturbances that generate canopy coverage gradients surrounding aquatic ecosystems is behavioural. The magnitude of the colonisation responses reported here rivals, if not exceeds, those produced by predators, suggesting that aquatic landscapes are behaviourally assessed and partitioned across multiple environmental gradients. The community level structure produced solely by selective colonisation, is predicted to strongly modify how patch area and isolation affect colonisation rates and the degree to which communities are linked by the flux of individuals and species
Metabolic Rate Models and the Substitutability of Predator Populations
1. Much of the debate surrounding the consequences of biodiversity loss centres around the issue of whether different species are functionally similar in their effects on ecological processes. In this study, we examined whether populations consisting of smaller, more abundant individuals are functionally similar to populations of the same species with larger, fewer individuals. 2. We manipulated the biomass and density of banded sunfish (Enneacanthus obesus) and measured their impact on populations of Southern leopard frog (Rana sphenocephala) larvae. We also evaluated the ability of models relating metabolic rate to body size to predict the relative impacts of populations that differ in average body size and population density. 3. Our results indicate that population biomass, density and their interaction each play a large role in determining the effect of a predator population on its food resource. Populations with smaller but more abundant individuals had effects as large or larger than those populations with larger but fewer individuals. 4. Although we found qualitative agreement between the observed relative effects of populations with that predicted by allometric models, we also found that density-dependence can cause effects of a population to differ from that expected based on allometry. 5. The substitutability of populations differing in average body size appears to depend on complex relationships between metabolic rate, population density and the strength of density-dependence. The restrictive conditions necessary to establish functional equivalence among different populations of the same species suggests that functional equivalence should be rare in most communities
Mismatched anti-predator behavioral responses in predator-naïve larval anurans
Organisms are adept at altering behaviors to balance the tradeoff between foraging and predation risk in spatially and temporally shifting predator environments. In order to optimize this tradeoff, prey need to be able to display an appropriate response based on degree of predation risk. To be most beneficial in the earliest life stages in which many prey are vulnerable to predation, innate anti-predator responses should scale to match the risk imposed by predators until learned anti-predator responses can occur. We conducted an experiment that examined whether tadpoles with no previous exposure to predators (i.e., predator-naive) exhibit innate antipredator behavioral responses (e.g., via refuge use and spatial avoidance) that match the actual risk posed by each predator. Using 7 treatments (6 free-roaming, lethal predators plus no-predator control), we determined the predation rates of each predator on Lithobates sphenocephalus tadpoles. We recorded behavioral observations on an additional 7 nonlethal treatments (6 caged predators plus no-predator control). Tadpoles exhibited innate responses to fish predators, but not non-fish predators, even though two non-fish predators (newt and crayfish) consumed the most tadpoles. Due to a mismatch between innate response and predator consumption, tadpoles may be vulnerable to greater rates of predation at the earliest life stages before learning can occur. Thus, naïve tadpoles in nature may be at a high risk to predation in the presence of a novel predator until learned anti-predator responses provide additional defenses to the surviving tadpoles.ECU Open Access Publishing Support Fun
Oviposition Behavior Partitions Aquatic Landscapes Along Predation and Nutrient Gradients
That individuals attempt to minimize the ratio of mortality risk/growth rate (μ/g) when foraging within individual habitat patches is well established. Do species partition among spatially discrete communities embedded in complex landscapes in a similar manner? We investigated how 3 ovipositing species (2 Hyla treefrogs and a hydrophilid beetle, Tropisternus lateralis) responded to simultaneous gradients of nutrients and predation risk. Species partitioned our experimental metacommunity primarily by reducing oviposition with fish. Tropisternus positively responded to increased nutrients, but the effect decreased with increasing risk, as predicted by μ/g theory. Use of fish habitats by Tropisternus was unrelated to breeding intensity. In contrast, Hyla showed no nutrient response but oviposited with fish only on nights with high breeding activity. Behavioral responses to the spatial distribution of resources and risk among discrete patches generated substantial variation in habitat-specific colonization rates, which has been identified as a primary mechanism generating both community and metacommunity structure
Temperature but not nutrient addition affects abundance and assemblage structure of colonizing aquatic insects
Abiotic conditions are important considerations in the species sorting process, which ultimately determines the distribution and abundance of species. Freshwater ecosystems will be impacted by ongoing temperature rise and other anthropogenically induced changes, such as nutrient enrichment and eutrophication. Changing characteristics of freshwater habitats will likely impact organisms in numerous ways, including through effects on colonization dynamics. Species are expected to colonize habitat patches where fitness will be the highest for themselves and their offspring, and how habitat selection interacts with changing environments remains an important question. We conducted a warming experiment to test the habitat selection preferences of aquatic beetles and hemipterans between habitat patches (mesocosms) of varying temperatures (via heaters), nutrient addition, and their interaction. Overall, insect abundance and richness were higher in unheated patches, with taxon-specific variation in response to heating. Although nutrients had limited effects on environmental conditions in mesocosms, their addition had no significant effects on insects. Insect assemblages had unique structures across heating treatments, with lower beta diversity and higher effective numbers of species in the warmest mesocosms. Our data support the importance of spatial variation in abiotic factors during the habitat selection process, and in determining species distributions and abundances as shallow lentic ecosystems are impacted by rising global temperatures
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