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AbstractDetermining whether and how evolution is predictable is an important goal, particularly as anthropogenic disturbances lead to novel species interactions that could modify selective pressures. Here, we use a multigeneration field experiment with brown anole lizards (Anolis sagrei) to test hypotheses about the predictability of evolution. We manipulated the presence/absence of predators and competitors of A. sagrei across 16 islands in the Bahamas that had preexisting brown anole populations. Before the experiment and again after roughly five generations, we measured traits related to locomotor performance and habitat use by brown anoles and used double-digest restriction enzyme-associated DNA sequencing to estimate genome-wide changes in allele frequencies. Although previous work showed that predators and competitors had characteristic effects on brown anole behavior, diet, and population sizes, we found that evolutionary change at both phenotypic and genomic levels was difficult to forecast. Phenotypic changes were contingent on sex and habitat use, whereas genetic change was unpredictable and not measurably correlated with phenotypic changes, experimental treatments, or other environmental factors. Our work shows how differences in ecological context can alter evolutionary outcomes over short timescales and underscores the difficulty of forecasting evolutionary responses to multispecies interactions in natural conditions, even in a well-studied system with ample supporting ecological information.

Assuntos

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AbstractIncreases in consumer abundance following a resource pulse can be driven by diet shifts, aggregation, and reproductive responses, with combined responses expected to result in faster response times and larger numerical increases. Previous work in plots on large Bahamian islands has shown that lizards (Anolis sagrei) increased in abundance following pulses of seaweed deposition, which provide additional prey (i.e., seaweed detritivores). Numerical responses were associated with rapid diet shifts and aggregation, followed by increased reproduction. These dynamics are likely different on isolated small islands, where lizards cannot readily immigrate or emigrate. To test this, we manipulated the frequency and magnitude of seaweed resource pulses on whole small islands and in plots within large islands, and we monitored lizard diet and numerical responses over 4 years. We found that seaweed addition caused persistent increases in lizard abundance on small islands regardless of pulse frequency or magnitude. Increased abundance may have occurred because the initial pulse facilitated population establishment, possibly via enhanced overwinter survival. In contrast with a previous experiment, we did not detect numerical responses in plots on large islands, despite lizards consuming more marine resources in subsidized plots. This lack of a numerical response may be due to rapid aggregation followed by disaggregation or to stronger suppression of A. sagrei by their predators on the large islands in this study. Our results highlight the importance of habitat connectivity in governing ecological responses to resource pulses and suggest that disaggregation and changes in survivorship may be underappreciated drivers of pulse-associated dynamics.

Assuntos

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Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect-in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence.

Assuntos

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Understanding processes that may stabilize ecological systems confronted with rapidly changing environmental conditions is a key issue in ecology. We studied a system of highly fluctuating populations, the moth Achyra rantalis feeding on the plant Sesuvium portulacastrum in a group of small subtropical islands of the Bahamas. The plant is a prostrate inhabitant of shorelines, and consequently moths are highly vulnerable to being consumed by the ground-foraging lizard Anolis sagrei. We measured the percent ground cover of Sesuvium and abundance of Achyra on 11 islands with lizards present and 21 islands without lizards annually for 10 consecutive years. Overall abundance of Achyra was 4.6 times higher on no-lizard islands than on lizard islands. The percent cover of Sesuvium exhibited lower temporal variability on lizard islands when the study site was undisturbed by hurricanes, and higher recovery rate on lizard islands following hurricanes. We suggest that both of these stabilizing phenomena are linked to a trophic cascade in which predatory lizards control herbivore populations, thereby suppressing outbreaks and enhancing plant recovery following physical disturbance.

Assuntos

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Flows of energy and materials link ecosystems worldwide and have important consequences for the structure of ecological communities. While these resource subsidies typically enter recipient food webs through multiple channels, most previous studies focussed on a single pathway of resource input. We used path analysis to evaluate multiple pathways connecting chronic marine resource inputs (in the form of seaweed deposits) and herbivory in a shoreline terrestrial ecosystem. We found statistical support for a fertilization effect (seaweed increased foliar nitrogen content, leading to greater herbivory) and a lizard numerical response effect (seaweed increased lizard densities, leading to reduced herbivory), but not for a lizard diet-shift effect (seaweed increased the proportion of marine-derived prey in lizard diets, but lizard diet was not strongly associated with herbivory). Greater seaweed abundance was associated with greater herbivory, and the fertilization effect was larger than the combined lizard effects. Thus, the bottom-up, plant-mediated effect of fertilization on herbivory overshadowed the top-down effects of lizard predators. These results, from unmanipulated shoreline plots with persistent differences in chronic seaweed deposition, differ from those of a previous experimental study of the short-term effects of a pulse of seaweed deposition: while the increase in herbivory in response to chronic seaweed deposition was due to the fertilization effect, the short-term increase in herbivory in response to a pulse of seaweed deposition was due to the lizard diet-shift effect. This contrast highlights the importance of the temporal pattern of resource inputs in determining the mechanism of community response to resource subsidies.

Assuntos

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The extent to which random processes such as founder events contribute to evolutionary divergence is a long-standing controversy in evolutionary biology. To determine the respective contributions of founder effects and natural selection, we conducted an experiment in which brown anole (Anolis sagrei) lizard populations were established on seven small islands in the Bahamas, from male-female pairs randomly drawn from the same large-island source. These founding events generated significant among-island genetic and morphological differences that persisted throughout the course of the experiment despite all populations adapting in the predicted direction-shorter hindlimbs-in response to the narrower vegetation on the small islands. Thus, using a replicated experiment in nature, we showed that both founder effects and natural selection jointly determine trait values in these populations.

Assuntos

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The effect of environmental change on ecosystems is mediated by species interactions. Environmental change may remove or add species and shift life-history events, altering which species interact at a given time. However, environmental change may also reconfigure multispecies interactions when both species composition and phenology remain intact. In a Caribbean island system, a major manifestation of environmental change is seaweed deposition, which has been linked to eutrophication, overfishing, and hurricanes. Here, we show in a whole-island field experiment that without seaweed two predators--lizards and ants--had a substantially greater-than-additive effect on herbivory. When seaweed was added to mimic deposition by hurricanes, no interactive predator effect occurred. Thus environmental change can substantially restructure food-web interactions, complicating efforts to predict anthropogenic changes in ecosystem processes.

Assuntos

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Ecologists have long struggled to explain variation in food-chain length among natural ecosystems. Food-chain length is predicted to be shorter in ecosystems subjected to greater disturbance because longer chains are theoretically less resilient to perturbation. Moreover, food-chain length is expected to be longer in larger ecosystems because increasing ecosystem size increases species richness and stabilizes predator-prey interactions, or increases total resource availability. Here we test the roles of disturbance and ecosystem size in determining the food-chain length of terrestrial food webs on Bahamian islands. We found that disturbance affected the identity of top predators, but did not change food-chain length because alternative top predators occupied similar trophic positions. On the other hand, a 106 -fold increase in ecosystem size elevated food-chain length by one trophic level. We suggest that the effect of disturbance on food-chain length is weak when alternate top predators are trophic omnivores and have similar trophic positions. This and previous work in lakes suggest that ecosystem size may be a strong determinant of food-chain length in both aquatic and terrestrial ecosystems.

RESUMO

We investigated how temporal variation in rainfall influences the impact of lizards on spiders inhabiting small islands in Abaco, Bahamas. Annual censuses of web spiders were conducted on nine lizard islands and on eight no-lizard islands 1994-2003. Repeated-measures ANOVA showed that annual variation in spider density (time) and in the lizard effect on spider density (lizard x time) were both significant. Correlation coefficients between the lizard effect (ln ratio of no-lizard to lizard spider densities) and number of rainfall days were generally negative, and strengthened with length of the time period during which rainfall was measured prior to annual spider censuses. Spider density was also negatively correlated with rainfall days and strengthened with length of the prior time period. Longer time intervals included the hurricane season, suggesting that the strong negative correlations were linked to high rainfall years during which tropical storms impacted the region and reduced spider and lizard densities. Split-plot ANOVA showed that rainfall during the hurricane season had a significant effect on the lizard effect and on spider density. Results in this study are opposite to those found in our previous 10-year study (1981-1990) conducted in the Exuma Cays, a moderately xeric region of the Bahamas, where the relation between rainfall and the lizard effect on spider density was positive. Combined data from the Exuma and Abaco studies produce a unimodal relation between trophic interaction strength and rainfall; we suggest that the negative effect of storms associated with rainfall was paramount in the present study, whereas the positive bottom-up effect of rainfall prevailed in our previous study. We conclude that climatic variability has a major impact on the trophic interaction and suggest that a substantial change in precipitation in either direction may weaken the interaction significantly.

Assuntos

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Major abiotic disturbance can be an important factor influencing food-web dynamics, particularly in areas impacted by the recent increase in hurricane activity. We present a unique set of data on key food-web processes occurring on 10 small islands for three relatively calm years and then four subsequent years during which two hurricanes passed directly over the study site. Herbivory, as measured by leaf damage, was 3.2 times higher in the year after the first hurricane (2000) than in the previous year and was 1.7 times higher in the year after the second hurricane (2002) than in 2001. The effect of a top predator (the lizard, Anolis sagrei) on herbivory strengthened continuously after the first hurricane and overall was 2.4 times stronger during the disturbance period than before. Overall abundance of lizards was 30% lower during the disturbance period than before, and abundances of web spiders and hymenopteran parasitoids were 66% and 59% lower, respectively. We suggest that increased herbivory observed on all islands was caused, at least in part, by the overall reduction in predation by both lizards and arthropods, whereas magnification of the lizard effect on herbivory was caused by reduced compensatory predation by arthropods.

Assuntos

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As the environment changes, will species be able to adapt? By conducting experiments in natural environments, biologists can study how evolutionary processes such as natural selection operate through time. We predicted that the introduction of a terrestrial predator would first select for longer-legged lizards, which are faster, but as the lizards shifted onto high twigs to avoid the predator, selection would reverse toward favoring the shorter-legged individuals better able to locomote there. Our experimental studies on 12 islets confirmed these predictions within a single generation, thus demonstrating the rapidity with which evolutionary forces can change during times of environmental flux.

Assuntos

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We monitored spiders on 41 Bahamian islands for 4 years before and then 4 years after the catastrophic Hurricane Floyd passed directly over the site, inundating the study islands with its storm surge. The respective recoveries of major community properties after this annihilation were far from synchronous. Before the hurricane, the species-area relation was generally strong and the slope showed no temporal trend. After the hurricane, the slope increased from near zero (7 months later) to a value about equal to its prehurricane state. The lizard effect (difference in spider abundance or species richness between islands with and without the lizard Anolis sagrei) was generally strong before the hurricane; 7 months after, the lizard effect on abundance was weak and the effect on richness had vanished. In subsequent years, the lizard effect on abundance became strong again, but the effect on species richness remained weak. The strength of the lizard effect on both abundance and richness over the 8 years was strongly positively related to the density of lizards measured on a subset of the study islands. Twelve months after the hurricane, species richness averaged over all islands rebounded to the last prehurricane value, but abundance attained only about half that value; this finding was remarkably similar to results found in an earlier study of spiders impacted by Hurricane Lili (1996) in a different Bahamian region. Nonetheless, in the next 3 years, species richness failed to increase further, part of its long-term decline at the study site.

Assuntos

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Population phenomena, which provide much of the underlying basis for the theoretical structure of island biogeography, have received little direct study. We determined a key population trait-survival-in the Bahamian lizard Anolis sagrei on islands with an experimentally introduced predatory lizard and on neighboring unmanipulated islands. On unmanipulated islands, survival declined with several variables, most notably vegetation height: The island with the shortest vegetation had nearly the highest survival recorded for any lizard. On islands with the introduced predator, which forages mostly on the ground, A. sagrei shifted to taller vegetation; unlike on unmanipulated islands, its survival was very low on islands with the shortest vegetation but was higher on the others. Thus, species introduction radically changed a resident species' relation of survival to a key island-biogeographical variable.

Assuntos

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The role of behaviour in evolutionary change has long been debated. On the one hand, behavioural changes may expose individuals to new selective pressures by altering the way that organisms interact with the environment, thus driving evolutionary divergence. Alternatively, behaviour can act to retard evolutionary change: by altering behavioural patterns in the face of new environmental conditions, organisms can minimize exposure to new selective pressures. This constraining influence of behaviour has been put forward as an explanation for evolutionary stasis within lineages and niche conservatism within clades. Nonetheless, the hypothesis that behavioural change prevents natural selection from operating in new environments has never been experimentally tested. We conducted a controlled and replicated experimental study of selection in entirely natural populations; we demonstrate that lizards alter their habitat use in the presence of an introduced predator, but that these behavioural shifts do not prevent patterns of natural selection from changing in experimental populations.

Assuntos

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Two recent hurricanes passed directly over the northern Bahamas 2 years apart, allowing a comparison of their effects on lizard populations inhabiting exactly the same islands. The hurricanes differed in two ways: one struck during the reproductive season and was relatively severe; the other struck after most reproduction had taken place and was milder. The late-season hurricane produced a significant relation between population reduction and lowness of the island that lasted at least through two seasons; the earlier hurricane produced no such relationship. The late-season hurricane wiped out populations of lizards on two islands (two of the three lowest) that the earlier hurricane failed to exterminate even though it was stronger. We relate these effects to the fact that the study lizards regenerated from the earlier hurricane only via the egg stage, whereas eggs were unavailable when the later storm struck and regeneration was via hatched lizards. We discriminate and illustrate four kinds of hurricanes, cross-classified by two contrasts: earlier vs. later and stronger vs. weaker. A later, stronger hurricane completely exterminated lizard populations at a second Bahamian site, whereas an earlier, weaker hurricane had no detectable effect at a third Bahamian site. We suggest that, in addition to severity, the timing of a hurricane as it coincides with reproductive scheduling or other phenological aspects may determine the magnitude of its effect on a variety of organisms.

Assuntos