Hurricane disturbance drives trophic changes in neotropical mountain stream food webs.

Food webs are complex ecological networks that reveal species interactions and energy flow in ecosystems. Prevailing ecological knowledge on forested streams suggests that their food webs are based on allochthonous carbon, driven by a constant supply of organic matter from adjacent vegetation and limited primary production due to low light conditions. Extreme climatic disturbances can disrupt these natural ecosystem dynamics by altering resource availability, which leads to changes in food web structure and functioning. Here, we quantify the response of stream food webs to two major hurricanes (Irma and María, Category 5 and 4, respectively) that struck Puerto Rico in September 2017. Within two tropical forested streams (first and second order), we collected ecosystem and food web data 6 months prior to the hurricanes and 2, 9, and 18 months afterward. We assessed the structural (e.g., canopy) and hydrological (e.g., discharge) characteristics of the ecosystem and monitored changes in basal resources (i.e., algae, biofilm, and leaf litter), consumers (e.g., aquatic invertebrates, riparian consumers), and applied Layman's community-wide metrics using the isotopic composition of 13 C and 15 N. Continuous stream discharge measurements indicated that the hurricanes did not cause an extreme hydrological event. However, the sixfold increase in canopy openness and associated changes in litter input appeared to trigger an increase in primary production. These food webs were primarily based on terrestrially derived carbon before the hurricanes, but most taxa (including Atya and Xiphocaris shrimp, the consumers with highest biomass) shifted their food source to autochthonous carbon within 2 months of the hurricanes. We also found evidence that the hurricanes dramatically altered the structure of the food web, resulting in shorter (i.e., smaller food-chain length), narrower (i.e., lower diversity of carbon sources) food webs, as well as increased trophic species packing. This study demonstrates how hurricane disturbance can alter stream food webs, changing the trophic base from allochthonous to autochthonous resources via changes in the physical environment (i.e., canopy defoliation). As hurricanes become more frequent and severe due to climate change, our findings greatly contribute to our understanding of the mechanisms that maintain forested stream trophic interactions amidst global change.

Landscape patterns in top-down control of decomposition: omnivory disrupts a tropical detrital-based trophic cascade.

Detrital-based trophic cascades are often considered weak or absent in tropical stream ecosystems because of the prevalence of omnivorous macroconsumers and the dearth of leaf-shredding insects. In this study, we isolate top-down effects of three macroconsumer species on detrital processing in headwater streams draining Trinidad's northern mountains. We separated effects of different macroconsumers by experimentally manipulating their temporal access to isolated benthic habitat over the diel cycle. We found no evidence that omnivorous macroconsumers, including a freshwater crab (Eudaniela garmani) and guppy (Poecilia reticulata), increased leaf decomposition via consumption. By contrast, above a waterfall excluding guppies, the insectivorous killifish, Anablepsoides hartii, reduced the biomass of the leaf-shredding insect Phylloicus hansoni 4-fold, which consequently reduced leaf decomposition rates 1.6-fold. This detrital cascade did not occur below the barrier waterfall, where omnivorous guppies join the assemblage and reduce killifish densities; here killifish had no significant effects on Phylloicus or decomposition rates. These patterns of detrital processing were also observed in upstream-downstream comparisons in a landscape study across paired reaches of six streams. Above waterfalls, where killifish were present, but guppies absent, leaf decomposition rates and Phylloicus biomass were 2.5- and ~35-fold lower, respectively, compared to measurements below waterfalls. Moreover, the strength of top-down control by killifish is reflected by the 20- and 5-fold reductions in variability (±SE) surrounding mean Phylloicus biomass and leaf decomposition rates in upstream relative to downstream reaches where no top-down control was detected. Findings show a clear, detrital-based trophic cascade among killifish, a leaf-shredding insect, and leaf decomposition rates. Results also show how omnivorous guppies disrupt this cascade by depressing killifish densities, thereby releasing invertebrate shredders from predation, and significantly increasing decomposition rates. Moreover, this combination of direct and indirect trophic interactions drives patterns in decomposition rates in stream networks at a landscape scale, resulting in significantly lower rates of decomposition above vs. below barrier waterfalls. Our findings reveal that omnivory can result in significant indirect effects on a key ecosystem process, illustrating the importance of these hidden trophic pathways in detrital-based systems and suggesting that resource control in tropical systems may be even more complex than previously envisioned.

Building a hydrologic foundation for tropical watershed management.

Here we provide an empirical hydrologic foundation to inform water management decisions in the El Yunque National Forest (EYNF) in eastern Puerto Rico. Tropical watershed hydrology has proven difficult to quantify due to high rainfall variability, high evapotranspiration rates, variation in forest canopy interception and storage, and uncertain hydrologic inputs from fog condensation in cloud forests. We developed mass-balance and observation-based water budgets for nine local watersheds within the EYNF using a novel assemblage of remotely sensed rainfall data, gaged streamflow observations, and municipal water withdrawal rates. It is important to note that, while prior budgets considered large water withdrawals outside (downstream) of EYNF boundaries, our current budget is confined to within EYNF boundaries. Here, we also base our estimates of water withdrawal volume on operational data, in contrast to prior water budgets that estimated volume based on either the capacity of known water intakes or regulatory permit limits. This resulted in more conservative and realistic estimates of withdrawals from within the EYNF. Finally, we also discuss the ecological importance of considering the effects of water withdrawals not only at an average monthly scale, but also on the basis of exceedance probability to avoid over-abstraction for the protection of native migratory fishes and shrimps. This analysis highlights a number of unique challenges associated with developing hydrologic foundations for water management in tropical ecosystems.

Large-scale climatic phenomena drive fluctuations in macroinvertebrate assemblages in lowland tropical streams, Costa Rica: The importance of ENSO events in determining long-term (15y) patterns.

Understanding how environmental variables influence the distribution and density of organisms over relatively long temporal scales is a central question in ecology given increased climatic variability (e.g., precipitation, ENSO events). The primary goal of our study was to evaluate long-term (15y time span) patterns of climate, as well as environmental parameters in two Neotropical streams in lowland Costa Rica, to assess potential effects on aquatic macroinvertebrates. We also examined the relative effects of an 8y whole-stream P-enrichment experiment on macroinvertebrate assemblages against the backdrop of this long-term study. Climate, environmental variables and macroinvertebrate samples were measured monthly for 7y and then quarterly for an additional 8y in each stream. Temporal patterns in climatic and environmental variables showed high variability over time, without clear inter-annual or intra-annual patterns. Macroinvertebrate richness and abundance decreased with increasing discharge and was positively related to the number of days since the last high discharge event. Findings show that fluctuations in stream physicochemistry and macroinvertebrate assemblage structure are ultimately the result of large-scale climatic phenomena, such as ENSO events, while the 8y P-enrichment did not appear to affect macroinvertebrates. Our study demonstrates that Neotropical lowland streams are highly dynamic and not as stable as is commonly presumed, with high intra- and inter-annual variability in environmental parameters that change the structure and composition of freshwater macroinvertebrate assemblages.

Riparian plant litter quality increases with latitude.

Plant litter represents a major basal resource in streams, where its decomposition is partly regulated by litter traits. Litter-trait variation may determine the latitudinal gradient in decomposition in streams, which is mainly microbial in the tropics and detritivore-mediated at high latitudes. However, this hypothesis remains untested, as we lack information on large-scale trait variation for riparian litter. Variation cannot easily be inferred from existing leaf-trait databases, since nutrient resorption can cause traits of litter and green leaves to diverge. Here we present the first global-scale assessment of riparian litter quality by determining latitudinal variation (spanning 107°) in litter traits (nutrient concentrations; physical and chemical defences) of 151 species from 24 regions and their relationships with environmental factors and phylogeny. We hypothesized that litter quality would increase with latitude (despite variation within regions) and traits would be correlated to produce 'syndromes' resulting from phylogeny and environmental variation. We found lower litter quality and higher nitrogen:phosphorus ratios in the tropics. Traits were linked but showed no phylogenetic signal, suggesting that syndromes were environmentally determined. Poorer litter quality and greater phosphorus limitation towards the equator may restrict detritivore-mediated decomposition, contributing to the predominance of microbial decomposers in tropical streams.

Population variation in the trophic niche of the Trinidadian guppy from different predation regimes.

Population variation in trophic niche is widespread among organisms and is of increasing interest given its role in both speciation and adaptation to changing environments. Trinidadian guppies (Poecilia reticulata) inhabiting stream reaches with different predation regimes have rapidly evolved divergent life history traits. Here, we investigated the effects of both predation and resource availability on guppy trophic niches by evaluating their gut contents, resource standing stocks, and δ15N and δ13C stable isotopes across five streams during the wet season. We found that guppies from low predation (LP) sites had a consistently higher trophic position and proportion of invertebrates in their guts and assimilate less epilithon than guppies from high predation (HP) sites. Higher trophic position was also associated with lower benthic invertebrate availability. Our results suggest that LP guppies could be more efficient invertebrate consumers, possibly as an evolutionary response to greater intraspecific competition for higher quality food. This may be intensified by seasonality, as wet season conditions can alter resource availability, feeding rates, and the intensity of intraspecific competition. Understanding how guppy diets vary among communities is critical to elucidating the role of niche shifts in mediating the link between environmental change and the evolution of life histories.

Disassembly of a tadpole community by a multi-host fungal pathogen with limited evidence of recovery.

Emerging infectious diseases can cause host community disassembly, but the mechanisms driving the order of species declines and extirpations following a disease outbreak are unclear. We documented the community disassembly of a Neotropical tadpole community during a chytridiomycosis outbreak, triggered by the generalist fungal pathogen, Batrachochytrium dendrobatidis (Bd). Within the first 11 months of Bd arrival, tadpole density and occupancy rapidly declined. Species rarity, in terms of tadpole occupancy and adult relative abundance, did not predict the odds of tadpole occupancy declines. But species losses were taxonomically selective, with glassfrogs (Family: Centrolenidae) disappearing the fastest and tree frogs (Family: Hylidae) and dart-poison frogs (Family: Dendrobatidae) remaining the longest. We detected biotic homogenization of tadpole communities, with post-decline communities resembling one another more strongly than pre-decline communities. The entire tadpole community was extirpated within 22 months following Bd arrival, and we found limited signs of recovery within 10 years post-outbreak. Because of imperfect species detection inherent to sampling species-rich tropical communities and the difficulty of devising a single study design protocol to sample physically complex tropical habitats, we used simulations to provide recommendations for future surveys to adequately sample diverse Neotropical communities. Our unique data set on tadpole community composition before and after Bd arrival is a valuable baseline for assessing amphibian recovery. Our results are of direct relevance to conservation managers and community ecologists interested in understanding the timing, magnitude, and consequences of disease outbreaks as emerging infectious diseases spread globally.

Fish introductions and light modulate food web fluxes in tropical streams: a whole-ecosystem experimental approach.

Decades of ecological study have demonstrated the importance of top-down and bottom-up controls on food webs, yet few studies within this context have quantified the magnitude of energy and material fluxes at the whole-ecosystem scale. We examined top-down and bottom-up effects on food web fluxes using a field experiment that manipulated the presence of a consumer, the Trinidadian guppy Poecilia reticulata, and the production of basal resources by thinning the riparian forest canopy to increase incident light. To gauge the effects of these reach-scale manipulations on food web fluxes, we used a nitrogen (15 N) stable isotope tracer to compare basal resource treatments (thinned canopy vs. control) and consumer treatments (guppy introduction vs. control). The thinned canopy stream had higher primary production than the natural canopy control, leading to increased N fluxes to invertebrates that feed on benthic biofilms (grazers), fine benthic organic matter (collector-gatherers), and organic particles suspended in the water column (filter feeders). Stream reaches with guppies also had higher primary productivity and higher N fluxes to grazers and filter feeders. In contrast, N fluxes to collector-gatherers were reduced in guppy introduction reaches relative to upstream controls. N fluxes to leaf-shredding invertebrates, predatory invertebrates, and the other fish species present (Hart's killifish, Anablepsoides hartii) did not differ across light or guppy treatments, suggesting that effects on detritus-based linkages and upper trophic levels were not as strong. Effect sizes of guppy and canopy treatments on N flux rates were similar for most taxa, though guppy effects were the strongest for filter feeding invertebrates while canopy effects were the strongest for collector-gatherer invertebrates. Combined, these results extend previous knowledge about top-down and bottom-up controls on ecosystems by providing experimental, reach-scale evidence that both pathways can act simultaneously and have equally strong influence on nutrient fluxes from inorganic pools through primary consumers.

Biotic and abiotic variables influencing plant litter breakdown in streams: a global study.

Plant litter breakdown is a key ecological process in terrestrial and freshwater ecosystems. Streams and rivers, in particular, contribute substantially to global carbon fluxes. However, there is little information available on the relative roles of different drivers of plant litter breakdown in fresh waters, particularly at large scales. We present a global-scale study of litter breakdown in streams to compare the roles of biotic, climatic and other environmental factors on breakdown rates. We conducted an experiment in 24 streams encompassing latitudes from 47.8° N to 42.8° S, using litter mixtures of local species differing in quality and phylogenetic diversity (PD), and alder (Alnus glutinosa) to control for variation in litter traits. Our models revealed that breakdown of alder was driven by climate, with some influence of pH, whereas variation in breakdown of litter mixtures was explained mainly by litter quality and PD. Effects of litter quality and PD and stream pH were more positive at higher temperatures, indicating that different mechanisms may operate at different latitudes. These results reflect global variability caused by multiple factors, but unexplained variance points to the need for expanded global-scale comparisons.

Evidence for the persistence of food web structure after amphibian extirpation in a Neotropical stream.

Species losses are predicted to simplify food web structure, and disease-driven amphibian declines in Central America offer an opportunity to test this prediction. Assessment of insect community composition, combined with gut content analyses, was used to generate periphyton-insect food webs for a Panamanian stream, both pre- and post-amphibian decline. We then used network analysis to assess the effects of amphibian declines on food web structure. Although 48% of consumer taxa, including many insect taxa, were lost between pre- and post-amphibian decline sampling dates, connectance declined by less than 3%. We then quantified the resilience of food web structure by calculating the number of expected cascading extirpations from the loss of tadpoles. This analysis showed the expected effects of species loss on connectance and linkage density to be more than 60% and 40%, respectively, than were actually observed. Instead, new trophic linkages in the post-decline food web reorganized the food web topology, changing the identity of "hub" taxa, and consequently reducing the effects of amphibian declines on many food web attributes. Resilience of food web attributes was driven by a combination of changes in consumer diets, particularly those of insect predators, as well as the appearance of generalist insect consumers, suggesting that food web structure is maintained by factors independent of the original trophic linkages.

Experimental acidification of two biogeochemically-distinct neotropical streams: buffering mechanisms and macroinvertebrate drift.

Research into the buffering mechanisms and ecological consequences of acidification in tropical streams is lacking. We have documented seasonal and episodic acidification events in streams draining La Selva Biological Station, Costa Rica. Across this forested landscape, the severity in seasonal and episodic acidification events varies due to interbasin groundwater flow (IGF). Streams that receive IGF have higher concentrations of solutes and more stable pH (~6) than streams that do not receive IGF (pH ~5). To examine the buffering capacity and vulnerability of macroinvertebrates to short-term acidification events, we added hydrochloric acid to acidify a low-solute, poorly buffered (without IGF) and a high-solute, well buffered stream (with IGF). We hypothesized that: 1) protonation of bicarbonate (HCO(3)(-)) would neutralize most of the acid added in the high-solute stream, while base cation release from the sediments would be the most important buffering mechanism in the low-solute stream; 2) pH declines would mobilize inorganic aluminum (Ali) from sediments in both streams; and 3) pH declines would increase macroinvertebrate drift in both streams. We found that the high-solute stream neutralized 745 µeq/L (96% of the acid added), while the solute poor stream only neutralized 27.4 µeq/L (40%). Protonation of HCO(3)(-) was an important buffering mechanism in both streams. Base cation, Fe(2+), and Ali release from sediments and protonation of organic acids also provided buffering in the low-solute stream. We measured low concentrations of Ali release in both streams (2-9 µeq/L) in response to acidification, but the low-solute stream released double the amount Ali per 100 µeq of acid added than the high solute stream. Macroinvertebrate drift increased in both streams in response to acidification and was dominated by Ephemeroptera and Chironomidae. Our results elucidate the different buffering mechanisms in tropical streams and suggest that low-solute poorly buffered streams might be particularly vulnerable to episodic acidification.

Direct and indirect ecosystem effects of evolutionary adaptation in the Trinidadian guppy (Poecilia reticulata).

Ecological and evolutionary processes may interact on the same timescale, but we are just beginning to understand how. Several studies have examined the net effects of adaptive evolution on ecosystem properties. However, we do not know whether these effects are confined to direct interactions or whether they propagate further through indirect ecological pathways. Even less well understood is how the combination of direct and indirect ecological effects of the phenotype promotes or inhibits evolutionary change. We coupled mesocosm experiments and ecosystem modeling to evaluate the ecological effects of local adaptation in Trinidadian guppies (Poecilia reticulata). The experiments show that guppies adapted to life with and without predators alter the ecosystem directly through differences in diet. The ecosystem model reveals that the small total indirect effect of the phenotype observed in the experiments is likely a combination of several large indirect effects that act in opposing directions. The model further suggests that these indirect effects can reverse the direction of selection that direct effects alone exert back on phenotypic variation. We conclude that phenotypic divergence can have major effects deep in the web of indirect ecological interactions and that even small total indirect effects can radically change the dynamics of adaptation.

Global distribution of a key trophic guild contrasts with common latitudinal diversity patterns.

Most hypotheses explaining the general gradient of higher diversity toward the equator are implicit or explicit about greater species packing in the tropics. However, global patterns of diversity within guilds, including trophic guilds (i.e., groups of organisms that use similar food resources), are poorly known. We explored global diversity patterns of a key trophic guild in stream ecosystems, the detritivore shredders. This was motivated by the fundamental ecological role of shredders as decomposers of leaf litter and by some records pointing to low shredder diversity and abundance in the tropics, which contrasts with diversity patterns of most major taxa for which broad-scale latitudinal patterns haven been examined. Given this evidence, we hypothesized that shredders are more abundant and diverse in temperate than in tropical streams, and that this pattern is related to the higher temperatures and lower availability of high-quality leaf litter in the tropics. Our comprehensive global survey (129 stream sites from 14 regions on six continents) corroborated the expected latitudinal pattern and showed that shredder distribution (abundance, diversity and assemblage composition) was explained by a combination of factors, including water temperature (some taxa were restricted to cool waters) and biogeography (some taxa were more diverse in particular biogeographic realms). In contrast to our hypothesis, shredder diversity was unrelated to leaf toughness, but it was inversely related to litter diversity. Our findings markedly contrast with global trends of diversity for most taxa, and with the general rule of higher consumer diversity at higher levels of resource diversity. Moreover, they highlight the emerging role of temperature in understanding global patterns of diversity, which is of great relevance in the face of projected global warming.

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams.

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.

Geostatistical modeling of the spatial distribution of sediment oxygen demand within a Coastal Plain blackwater watershed.

Blackwater streams are found throughout the Coastal Plain of the southeastern United States and are characterized by a series of instream floodplain swamps that play a critical role in determining the water quality of these systems. Within the state of Georgia, many of these streams are listed in violation of the state's dissolved oxygen (DO) standard. Previous work has shown that sediment oxygen demand (SOD) is elevated in instream floodplain swamps and due to these areas of intense oxygen demand, these locations play a major role in determining the oxygen balance of the watershed as a whole. This work also showed SOD rates to be positively correlated with the concentration of total organic carbon. This study builds on previous work by using geostatistics and Sequential Gaussian Simulation to investigate the patchiness and distribution of total organic carbon (TOC) at the reach scale. This was achieved by interpolating TOC observations and simulated SOD rates based on a linear regression. Additionally, this study identifies areas within the stream system prone to high SOD at representative 3rd and 5th order locations. Results show that SOD was spatially correlated with the differences in distribution of TOC at both locations and that these differences in distribution are likely a result of the differing hydrologic regime and watershed position. Mapping of floodplain soils at the watershed scale shows that areas of organic sediment are widespread and become more prevalent in higher order streams. DO dynamics within blackwater systems are a complicated mix of natural and anthropogenic influences, but this paper illustrates the importance of instream swamps in enhancing SOD at the watershed scale. Moreover, our study illustrates the influence of instream swamps on oxygen demand while providing support that many of these systems are naturally low in DO.

Local adaptation in Trinidadian guppies alters ecosystem processes.

Theory suggests evolutionary change can significantly influence and act in tandem with ecological forces via ecological-evolutionary feedbacks. This theory assumes that significant evolutionary change occurs over ecologically relevant timescales and that phenotypes have differential effects on the environment. Here we test the hypothesis that local adaptation causes ecosystem structure and function to diverge. We demonstrate that populations of Trinidadian guppies (Poecilia reticulata), characterized by differences in phenotypic and population-level traits, differ in their impact on ecosystem properties. We report results from a replicated, common garden mesocosm experiment and show that differences between guppy phenotypes result in the divergence of ecosystem structure (algal, invertebrate, and detrital standing stocks) and function (gross primary productivity, leaf decomposition rates, and nutrient flux). These phenotypic effects are further modified by effects of guppy density. We evaluated the generality of these effects by replicating the experiment using guppies derived from two independent origins of the phenotype. Finally, we tested the ability of multiple guppy traits to explain observed differences in the mesocosms. Our findings demonstrate that evolution can significantly affect both ecosystem structure and function. The ecosystem differences reported here are consistent with patterns observed across natural streams and argue that guppies play a significant role in shaping these ecosystems.

Deviation from strict homeostasis across multiple trophic levels in an invertebrate consumer assemblage exposed to high chronic phosphorus enrichment in a Neotropical stream.

A central tenet of ecological stoichiometry is that consumer elemental composition is relatively independent of food resource nutrient content. Although the P content of some invertebrate consumer taxa can increase as a consequence of P-enriched food resources, little is known about how ecosystem nutrient loading can affect the elemental composition of entire consumer assemblages. Here we examine the potential for P enrichment across invertebrate consumer assemblages in response to chronic high P loading. We measured elemental ratios in invertebrate consumers and basal food resources in a series of streams in lowland Costa Rica that range widely in P levels (2-135 microg l(-1) soluble reactive P). Streams with high P levels receive natural long-term (over millennia) inputs of solute-rich groundwater while low-P streams do not receive these solute-rich groundwater inputs. P content of leaf litter and epilithon increased fourfold across the natural P gradient, exceeding basal resource P content values reported in the literature from other nutrient-rich streams. Invertebrate consumers from the high-P study stream were elevated twofold in P content across multiple taxonomic and functional feeding groups, including predators. Our results strongly support the hypothesis that elevated P content in consumers feeding on P-enriched food resources is a consequence of deviation from strict homeostasis. In contrast to prior studies, we found that between-stream variation in P content of a given taxon greatly exceeded within-stream variation among different taxa, suggesting that environment may be as important as phylogeny in controlling consumer stoichiometry. Relaxing the assumption of strict homeostasis presents challenges and opportunities for advancing our understanding of how nutrient limitation affects consumer growth. Moreover, our findings may provide a window into the future of how chronic anthropogenic nutrient loading can alter stoichiometric relationships in food webs.

Experimental evaluation of evolution and coevolution as agents of ecosystem change in Trinidadian streams.

Evolution has been shown to be a critical determinant of ecological processes in some systems, but its importance relative to traditional ecological effects is not well known. In addition, almost nothing is known about the role of coevolution in shaping ecosystem function. Here, we experimentally evaluated the relative effects of species invasion (a traditional ecological effect), evolution and coevolution on ecosystem processes in Trinidadian streams. We manipulated the presence and population-of-origin of two common fish species, the guppy (Poecilia reticulata) and the killifish (Rivulus hartii). We measured epilithic algal biomass and accrual, aquatic invertebrate biomass, and detrital decomposition. Our results show that, for some ecosystem responses, the effects of evolution and coevolution were larger than the effects of species invasion. Guppy evolution in response to alternative predation regimes significantly influenced algal biomass and accrual rates. Guppies from a high-predation site caused an increase in algae relative to guppies from a low-predation site; algae effects were probably shaped by observed divergence in rates of nutrient excretion and algae consumption. Rivulus-guppy coevolution significantly influenced the biomass of aquatic invertebrates. Locally coevolved populations reduced invertebrate biomass relative to non-coevolved populations. These results challenge the general assumption that intraspecific diversity is a less critical determinant of ecosystem function than is interspecific diversity. Given existing evidence for contemporary evolution in these fish species, our findings suggest considerable potential for eco-evolutionary feedbacks to operate as populations adapt to natural or anthropogenic perturbations.

Phylogeography of an Island endemic, the Puerto Rican Freshwater Crab (Epilobocera sinuatifrons).

The endemic Puerto Rican crab, Epilobocera sinuatifrons (Pseudothelphusidae), has a freshwater-dependant life-history strategy, although the species has some capabilities for terrestrial movement as adults. In contrast to all other freshwater decapods on the island (e.g., caridean shrimp), E. sinuatifrons does not undertake amphidromous migration, and is restricted to purely freshwater habitats and adjacent riparian zones. As Puerto Rico has a dynamic geologic history, we predicted that both the life history of E. sinuatifrons and the geological history of the island would be important determinants of phylogeographic structuring in the species. Using a fragment of the cytochrome c oxidase subunit 1 mtDNA (mitochondrial DNA) gene, we tested for deviations from panmixia among and within rivers draining Puerto Rico and used statistical phylogeography to explore processes that may explain extant patterns of genetic variation in the species. While populations of E. sinuatifrons were significantly differentiated among rivers, they were likely to be recently derived because nested clade analysis (NCA) indicated evolutionarily recent restricted gene flow with isolation by distance (IBD) and contiguous range expansion at various spatial scales. Ongoing drainage rearrangements associated with faulting and land slippage were invoked as processes involved in sporadic gene flow among rivers throughout the Pleistocene. Patterns of genetic differentiation conformed to IBD and population demographic statistics were nonsignificant, indicating that although recently derived, populations from different rivers were in drift-mutation equilibrium. A shallow (0.6 million years ago), paraphyletic split was observed in the haplotype network, which NCA indicated arose via allopatric fragmentation. This split coincides with an area of high relief in central Puerto Rico that may have experienced relatively little drainage rearrangements. Shallow but significant genetic isolation of populations of E. sinuatifrons among Puerto Rican rivers suggests phylogeographic patterns that are intermediate to terrestrial habitat specialists (highly divergent populations) and other freshwater biota, such as amphidromous species and insects with aerial adult dispersal (highly connected populations).

Molecular evidence for sequential colonization and taxon cycling in freshwater decapod shrimps on a Caribbean island.

Taxon cycling, i.e. sequential phases of expansions and contractions in species' distributions associated with ecological or morphological shifts, are postulated to characterize dynamic biogeographic histories in various island faunas. The Caribbean freshwater shrimp assemblage is mostly widespread and sympatric throughout the region, although one species (Atyidae: Atya lanipes) is geographically restricted and ecologically and morphologically differentiated from other Atya species. Using patterns of nucleotide variation at the COI mtDNA gene in five species of freshwater shrimp (A. lanipes, A. scabra, A. innocuous; Xiphocarididae: Xiphocaris elongata; Palaemonidae: Macrobrachium faustinum) from Puerto Rico, we expected to detect a signature of sequential colonization in these shrimp, consistent with the concept of taxon cycling, and expected that A. lanipes would be at a different taxon stage (i.e. an early stage species) to all other species. We also examined patterns of genetic population structure in each species expected with poor, intermediate and well-developed abilities for among-river dispersal. Population expansions were detected in all species, although the relative timing of the expansions varied among them. Assuming that population expansions followed colonization of Puerto Rico by freshwater shrimp, results bear the hallmarks of sequential colonization and taxon cycling in this fauna. A. lanipes had a star phylogeny, low mean pairwise nucleotide differences and recent (Holocene) estimates for an in situ population expansion in Puerto Rico, and it was inferred as an early stage species in the taxon cycle undergoing a secondary phase of expansion. All other species were inferred as late stage species undergoing regional population expansions, as their mean pairwise nucleotide differences were relatively high and phylogenetic patterns were more complex than A. lanipes. High rates of gene flow without isolation by distance among rivers were detected in all species, although results should be treated cautiously as some populations are unlikely to be in mutation-drift equilibrium. Nested clade analysis produced inconsistent results among species that all have high rates of gene flow and expanding populations.