Eugenia Zandonà

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.

The costs of hemispheric specialization in a fish

Laboratory and field studies have documented better cognitive performance associated with marked hemispheric specialization in organisms as diverse as chimpanzees, domestic chicks and topminnows. While providing an evolutionary explanation for the emergence of cerebral lateralization, this evidence represents a paradox because a large proportion of non-lateralized (NL) individuals is commonly observed in animal populations. Hemispheric specialization often determines large left–right differences in perceiving and responding to stimuli. Using topminnows selected for a high or low degree of lateralization, we tested the hypothesis that individuals with greater functional asymmetry pay a higher performance cost in situations requiring matching information from the two eyes. When trained to use the middle door in a row of a nine, NL fish correctly chose the central door in most cases, while lateralized fish showed systematic leftward or rightward biases. When choosing between two shoals, each seen with a different eye, NL fish chose the high-quality shoal significantly more often than the lateralized fish, whose performance was affected by eye preference for analysing social stimuli. These findings suggest the existence of a trade-off between computational advantages of hemispheric specialization and the ecological cost of making suboptimal decisions whenever relevant information is located on both sides of the body.

Flow, nutrients, and light availability influence Neotropical epilithon biomass and stoichiometry

Abstract.  Light, nutrient availability, and flow are strong factors controlling the elemental composition and biomass of epilithon in temperate stream ecosystems. However, comparatively little is known about these relationships in tropical streams. We investigated how gradients of light and nutrient availability, seasonality, and habitat influenced epilithon biomass, chlorophyll a, and nutrient ratios in montane streams of Trinidad, West Indies. We sampled 4 focal tributaries of a single river, 2 of which had canopies experimentally thinned, every other month over a 2-y period to observe temporal dynamics and light effects on epilithon. We also sampled 18 sites across Trinidads Northern Range Mountains once each in a wet and dry season to examine the effects of naturally occurring differences in light and dissolved nutrient availability on epilithic characteristics. We found greater chlorophyll a concentrations in habitats with greater light availability, but the effect of light on epilithon stoichiometry differed between the site-survey and focal-tributary data. In general, epilithic C∶nutrient ratios decreased with increasing dissolved nutrient concentrations, but relationships between nutrient availability and biomass probably were obscured by naturally high dissolved N and P concentrations in many of the streams. Season and habitat type had profound effects on epilithon variables. Biomass and % C generally decreased in riffles and under wet-season conditions. These results suggest multiple controls for the quantity and quality of stream epilithon and have important implications for in-stream consumers.

Environmental and organismal predictors of intraspecific variation in the stoichiometry of a neotropical freshwater fish.

The elemental composition of animals, or their organismal stoichiometry, is thought to constrain their contribution to nutrient recycling, their interactions with other animals, and their demographic rates. Factors that affect organismal stoichiometry are generally poorly understood, but likely reflect elemental investments in morphological features and life history traits, acting in concert with the environmental availability of elements. We assessed the relative contribution of organismal traits and environmental variability to the stoichiometry of an insectivorous Neotropical stream fish, Rivulus hartii. We characterized the influence of body size, life history phenotype, stage of maturity, and environmental variability on organismal stoichiometry in 6 streams that differ in a broad suite of environmental variables. The elemental composition of R. hartii was variable, and overlapped with the wide range of elemental composition documented across freshwater fish taxa. Average %P composition was ∼3.2%(±0.6), average %N∼10.7%(±0.9), and average %C∼41.7%(±3.1). Streams were the strongest predictor of organismal stoichiometry, and explained up to 18% of the overall variance. This effect appeared to be largely explained by variability in quality of basal resources such as epilithon N∶P and benthic organic matter C∶N, along with variability in invertebrate standing stocks, an important food source for R. hartii. Organismal traits were weak predictors of organismal stoichiometry in this species, explaining when combined up to 7% of the overall variance in stoichiometry. Body size was significantly and positively correlated with %P, and negatively with N∶P, and C∶P, and life history phenotype was significantly correlated with %C, %P, C∶P and C∶N. Our study suggests that spatial variability in elemental availability is more strongly correlated with organismal stoichiometry than organismal traits, and suggests that the stoichiometry of carnivores may not be completely buffered from environmental variability. We discuss the relevance of these findings to ecological stoichiometry theory.

Challenges for interpreting stable isotope fractionation of carbon and nitrogen in tropical aquatic ecosystems

Summary Stable isotopes are useful for elucidating food webs, and oneessentialaspectofinterpretationisaccuratelydeterminingtheenrichment between trophic levels, especially when used inmixing models. The fractionation of the stable isotopes ofnitrogen(Δδ 15 N)andcarbon(Δδ 3 C)betweentrophiclevelsintropicalaquaticecosystemsseemstodiffercomparedtotypicalvalues found in temperate aquatic ecosystems of about 3.4‰forδ 15 Nand0.5‰forδ 13 C.Inrecentstudiesofuplandstreamecosystems in Panama, with and without tadpoles, we foundlower fractionation of δ 15 N, typically 1.0–1.7‰, and muchhigherfractionationofδ 13 C,typically1–1.6‰.Similarvalueshavebeenobservedinothertropicalsystems,includingPuertoRicoandCostaRicastreamsandLakeTanganyika.Theselargedifferencesintrophicfractionationareenigmatic.Weexploresources of variation in fractionation such as tissue turnoverrate,streamflow,taxonomicdifferences,functionalgroupdif-ferences, dietary balance, growth rate, and microbial activityto assess possible contributing factors. The ecological infor-mation embedded in this variation in trophic fractionationshould be embraced and exploited.

Emotional responsiveness in fish from lines artificially selected for a high or low degree of laterality

Evidence showing that cerebral asymmetries exist in a wide range of animals has prompted investigation into the advantages and disadvantages of brain lateralization. In the teleost fish Girardinus falcatus individuals selected for a high degree of lateralization (LAT) performed better than those fish selected for reduced lateralization (NL) in several tasks, including schooling, foraging and spatial orientation. These findings were interpreted as evidence of hemispheric specialization allowing more efficient parallel processing and thus better cognitive performance under conditions that require multitasking, but the possibility that the results may simply reflect line differences in behavioral/physiological coping styles (i.e. in their emotional responsiveness during the tests) could not be ruled out. To test the hypothesis that NL and LAT fish differ in coping style, the present study examined differences in response in these lines to a novel situation in four different conditions. NL and LAT fish did not differ in a behavioral measure of emotional response: their readiness to explore a new environment. After being isolated in a tight space they showed a similar increase in opercular beating rates, suggesting that their physiological response to an acute stressor was comparable. The overall tendency to remain close to a shoalmate after being moved to an unfamiliar place was similar in the two groups but a significant difference was found in the temporal pattern; LAT fish swam closer than NL to their mirror image in the initial stages but this difference was later reversed. NL and LAT males placed in a new, unfamiliar environment did not differ in the number of sexual acts performed but LAT males resumed sexual behavior earlier signifying that cerebral lateralization has some influence on the trade-off between predator surveillance and mating behavior. Although this study found some differences between NL and LAT lines in their response to novelty, present evidence does not seem sufficient to justify the rejection of the hypothesis that the better scores in complex tasks shown by LAT fish in previous studies were primarily due to a cognitive advantage associated with cerebral specialization.

Effects of Consumer Interactions on Benthic Resources and Ecosystem Processes in a Neotropical Stream

The effect of consumers on their resources has been demonstrated in many systems but is often confounded by trophic interactions with other consumers. Consumers may also have behavioral and life history adaptations to each other and to co-occurring predators that may additionally modulate their particular roles in ecosystems. We experimentally excluded large consumers from tile periphyton, leaves and natural benthic substrata using submerged electrified frames in three stream reaches with overlapping consumer assemblages in Trinidad, West Indies. Concurrently, we assessed visits to (non-electrified) control frames by the three most common large consumers–primarily insectivorous killifish (Rivulus hartii), omnivorous guppies (Poecilia reticulata) and omnivorous crabs (Eudaniela garmani). Consumers caused the greatest decrease in final chlorophyll a biomass and accrual rates the most in the downstream reach containing all three focal consumers in the presence of fish predators. Consumers also caused the greatest increase in leaf decay rates in the upstream reach containing only killifish and crabs. In the downstream reach where guppies co-occur with predators, we found significantly lower benthic invertebrate biomass in control relative to exclosure treatments than the midstream reach where guppies occur in the absence of predators. These data suggest that differences in guppy foraging, potentially driven by differences in their life history phenotype, may affect ecosystem structure and processes as much as their presence or absence and that interactions among consumers may further mediate their effects in these stream ecosystems.

Population level “flipperedness” in the eastern Pacific leatherback turtle

Limb preference is a behavioral indicator of lateralized brain function that was recently elucidated experimentally in lower vertebrates. We assessed natural spontaneous limb use of nesting eastern Pacific leatherback turtles by recording which hindlimb flipper was extended overtop the cloaca to cover the egg chamber during oviposition. We found a population level right bias in 1889 observations of 361 individuals. This is the first report of a limb preference in Testudinata.

Contrasting Population and Diet Influences on Gut Length of an Omnivorous Tropical Fish, the Trinidadian Guppy ( Poecilia reticulata )

Phenotypic plasticity is advantageous for organisms that live in variable environments. The digestive system is particularly plastic, responding to changes in diet. Gut length is the result of a trade-off between maximum nutrient absorption and minimum cost for its maintenance and it can be influenced by diet and by evolutionary history. We assessed variation in gut length of Trinidadian guppies (Poecilia reticulata) as a function of diet, season, ontogeny, and local adaptation. Populations of guppies adapted to different predation levels have evolved different life history traits and have different diets. We sampled guppies from sites with low (LP) and high predation (HP) pressure in the Aripo and Guanapo Rivers in Trinidad. We collected fish during both the dry and wet season and assessed their diet and gut length. During the dry season, guppies from HP sites fed mostly on invertebrates, while guppies in the LP sites fed mainly on detritus. During the wet season, the diet of LP and HP populations became very similar. We did not find strong evidence of an ontogenetic diet shift. Gut length was negatively correlated with the proportion of invertebrates in diet across fish from all sites, supporting the hypothesis that guppy digestive systems adapt in length to changes in diet. Population of origin also had an effect on gut length, as HP and LP fish maintained different gut lengths even in the wet season, when their diets were very similar and individuals in both types of populations fed mostly on detritus. Thus, both environment and population of origin influenced guppies gut length, but population of origin seemed to have a stronger effect. Our study also showed that, even in omnivorous fish, gut length adapted to different diets, being more evident when the magnitude of difference between animal and plant material in the diet was very large.

Intraspecific variability modulates interspecific variability in animal organismal stoichiometry.

Interspecific differences in organismal stoichiometry (OS) have been documented in a wide range of animal taxa and are of significant interest for understanding evolutionary patterns in OS. In contrast, intraspecific variation in animal OS has generally been treated as analytical noise or random variation, even though available data suggest intraspecific variability in OS is widespread. Here, we assess how intraspecific variation in OS affects inferences about interspecific OS differences using two co-occurring Neotropical fishes: Poecilia reticulata and Rivulus hartii. A wide range of OS has been observed within both species and has been attributed to environmental differences among stream systems. We assess the contributions of species identity, stream system, and the interactions between stream and species to variability in N:P, C:P, and C:N. Because predation pressure can impact the foraging ecology and life-history traits of fishes, we compare predictors of OS between communities that include predators, and communities where predators are absent. We find that species identity is the strongest predictor of N:P, while stream or the interaction of stream and species contribute more to the overall variation in C:P and C:N. Interspecific differences in N:P, C:P, and C:N are therefore not consistent among streams. The relative contribution of stream or species to OS qualitatively changes between the two predation communities, but these differences do not have appreciable effects in interspecific patterns. We conclude that although species identity is a significant predictor of OS, intraspecific OS is sometimes sufficient to overwhelm or obfuscate interspecific differences in OS.