Testes mass in the livebearing fish Brachyrhaphis rhabdophora (Poeciliidae) varies hypoallometrically with body size but not between predation environments.

In this study, we considered potential causes of variation in testis size in the livebearing fish Brachyrhaphis rhabdophora. We evaluated variation in testes mass among individual males and among populations that occupy different selective environments. First, we predicted that small males should allocate more to testes mass than large males (i.e., hypoallometric pattern) based on a sperm competition argument. Second, based on life history theory and associated differences in mortality rates between populations that coexist with many fish predators and those with few predators, we predicted that males in high-predation environments should allocate more to testes mass than males in habitats with few predators. Our results showed that small males allocated proportionally more to testes mass than larger males (slope of testes mass to body mass was hypoallometric). However, there was no effect of predator environment on testes mass independent of body size differences. In this system, size-specific patterns of reproductive allocation in males (hypoallometry) differ from that seen in females (hyperallometry). Allocation to testes mass may respond to differences in mortality rate through selection on body size.

Mating and aggressive behaviour of Brachyrhaphis olomina (Cyprinodontiformes: Poeciliidae).

Despite the increasing interest in the use of intromittent male genitalia and coercive mating behaviour in poeciliids, detailed studies of the mating behaviour of most species in this family are lacking. We describe here the mating and aggressive behaviours of Brachyrhaphis olomina, and correlate them with the condition of the female's ovum and embryos (immature, mature and pregnant). B. olomina performed a wide range of aggressive (sidle spread, tail beating, coordinate) and mating behaviours (approximation, touch, lateral display, touch-lateral display). Some behaviours (e.g. tail beating) are shared with other poeciliids, but two sexual behaviours (touch and lateral display) and one aggressive (coordinate) behaviour may be unique to B. olomina and were not reported in a previous study. Differences in male behaviour when paired with a female with mature ovum (more mating displays, no agonistic movements) suggest that males detect the female's reproductive condition from some distance. The distinctive nature of mating behaviour in B. olomina highlights the importance of studying different species to have a better understanding of the evolution of mating and aggressive behaviours in poeciliids. Digital video images related to the article are available at http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo01a, http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo02a and http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo03a.

Predation environment affects boldness temperament of neotropical livebearers.

Behavioral traits of individuals are important phenotypes that potentially interact with many other traits, an understanding of which may illuminate the evolutionary forces affecting populations and species. Among the five axes of temperament is the propensity to behave boldly in the presence of a perceived risk. To determine the effect of different predatorial regimes on boldness and fearfulness, we assessed the behavior of individuals in a novel portable swim chamber (i.e., forced open-field test) by Brachyrhaphis rhabdophora (n = 633). We used an information theoretic framework to compare generalized (logistic) linear fixed-effects models of predatorial regime (predator-free [n = 6] and predator [n = 4] sites), sex, and standard length (SL). Fish from predator sites were much more fearful in the novel arena than fish from nonpredator sites. This varied by length, but not by sex. At 48 mm SL, fish from nonpredator sites were 4.9 times more likely to express bold behavior (ambulation) in the novel swim chamber as fish from predator sites. Probabilities of "ambulating" within the swim chamber increased with size for nonpredator sites and decreased with size for predator sites.

Divergent predation environment between two sister species of livebearing fishes (Cyprinodontiformes: Poeciliidae) predicts boldness, activity, and exploration behavior / Ambientes de depredación divergente para dos especies hermanas de peces vivíparos (Cyprinodontiformes: Poeciliidae) predicen el comportamiento de audacia, actividad y exploración

Abstract:Predators can influence a variety of prey traits, including behavior. Traits such as boldness, activity rate, and tendency to explore can all be shaped by predation risk. Our study examines the effects of predation on these behaviors by considering a natural system in which two sister species of livebearing fishes, Brachyrhaphis roseni and B. terrabensis, experience divergent predation environments. In February of 2013, we collected fish in the Río Chiriquí Nuevo drainage, Chiriquí, Panama, and conducted behavioral assays. Using open-field behavioral assays, we evaluated both juveniles and adults, and males and females, to determine if there were differences in behavior between ontogenetic stages or between sexes. We assessed boldness as 'time to emerge' from a shelter into a novel environment, and subsequently measured activity and exploration within that novel environment. We predicted that B. roseni (a species that co-occurs with predators) would be more bold, more active, and more prone to explore, than B. terrabensis (a species that does not co-occur with predators). In total, we tested 17 juveniles, 21 adult males, and 20 adult females of B. roseni, and 19 juveniles, 19 adult males, and 18 adult females of B. terrabensis. We collected all animals from streams in Chiriquí, Panama in February 2013, and tested them following a short acclimation period to laboratory conditions. As predicted, we found that predation environment was associated with several differences in behavior. Both adult and juvenile B. roseni were more active and more prone to explore than B. terrabensis. However, we found no differences in boldness in either adults or juveniles. We also found a significant interaction between 'sex' and 'species' as predictors of boldness and exploration, indicating that predation environment can affect behaviors of males and females differently in each species. Our work demonstrates the importance of considering sex and life history stage when evaluating the evolution of behavior. Rev. Biol. Trop. 65 (1): 267-277. Epub 2017 March 01.

ResumenLos depredadores pueden influenciar una variedad de rasgos de la presa, incluyendo el comportamiento. Los rasgos tales como la audacia, nivel de actividad, y la tendencia a explorar; pueden ser moldeados por el riesgo de depredación. Nuestro estudio examina los efectos de la depredación en el comportamiento al considerar un sistema natural en el que dos especies hermanas de peces vivíparos, Brachyrhaphis roseni y B. terrabensis, se presentan en ambientes de depredación divergente. En febrero 2013, recolectamos peces en el drenaje del Río Chiriquí Nuevo, Chiriquí, Panamá y llevamos a cabo ensayos de comportamiento. Al usar ensayos de comportamiento en campo abierto, se evaluó el comportamiento en juveniles y adultos, machos y hembras, para determinar si los patrones de divergencia diferían entre las etapas ontogenéticas o entre sexos. Se evaluó la audacia como "tiempo en salir" de un refugio a un ambiente nuevo, y posteriormente se midió la actividad y la exploración dentro de ese nuevo ambiente. Nosotros predijimos que B. roseni (una especie que se presenta con los depredadores) sería más audaz, activa y propensa a explorar que B. terrabensis (una especie que no se presenta con los depredadores). En total, probamos 17 jóvenes, 21 machos adultos, y 20 hembras adultas de B. roseni, y 19 jóvenes, 19 machos adultos, y 18 hembras adultas de B. terrabensis. Recogimos todos los animales en Chiriquí, Panamá en Febrero 2013, y los probamos después de un corto período de aclimatación a las condiciones de laboratorio. Como se predijo, se encontró que la depredación ambiental se asoció con varias diferencias en el comportamiento. Tanto adultos y jóvenes de B. roseni eran más activos y más propensos a explorar que B. terrabensis. Sin embargo, no se encontraron diferencias en la audacia en adultos o menores. También se encontró una interacción significativa entre "sexo" y "especie" como predictores de la audacia y la exploración, lo que indica que la depredación puede afectar el comportamiento de los machos y hembras de manera diferente en cada especie. Nuestro estudio demuestra la importancia de considerar el sexo y la etapa del ciclo de vida al evaluar la evolución del comportamiento.

Selection is stronger in early-versus-late stages of divergence in a Neotropical livebearing fish.

How selection acts to drive trait evolution at different stages of divergence is of fundamental importance in our understanding of the origins of biodiversity. Yet, most studies have focused on a single point along an evolutionary trajectory. Here, we provide a case study evaluating the strength of divergent selection acting on life-history traits at early-versus-late stages of divergence in Brachyrhaphis fishes. We find that the difference in selection is stronger in the early-diverged population than the late-diverged population, and that trait differences acquired early are maintained over time.

Divergent natural selection promotes immigrant inviability at early and late stages of evolutionary divergence.

Natural selection's role in speciation has been of fundamental importance since Darwin first outlined his theory. Recently, work has focused on understanding how selection drives trait divergence, and subsequently reproductive isolation. "Immigrant inviability," a barrier that arises from selection against immigrants in their nonnative environment, appears to be of particular importance. Although immigrant inviability is likely ubiquitous, we know relatively little about how selection acts on traits to drive immigrant inviability, and how important immigrant inviability is at early-versus-late stages of divergence. We present a study evaluating the role of predation in the evolution of immigrant inviability in recently diverged population pairs and a well-established species pair of Brachyrhaphis fishes. We evaluate performance in a high-predation environment by assessing survival in the presence of a predator, and swimming endurance in a low-predation environment. We find strong signatures of local adaptation and immigrant inviability of roughly the same magnitude both early and late in divergence. We find remarkably conserved selection for burst-speed swimming (important in predator evasion), and selection for increased size in low-predation environments. Our results highlight the consistency with which selection acts during speciation, and suggest that similar factors might promote initial population differentiation and maintain differentiation at late stages of divergence.

Phylogenetic analyses provide insights into the historical biogeography and evolution of Brachyrhaphis fishes.

The livebearing fish genus Brachyrhaphis (Poeciliidae) has become an increasingly important model in evolution and ecology research, yet the phylogeny of this group is not well understood, nor has it been examined thoroughly using modern phylogenetic methods. Here, we present the first comprehensive phylogenetic analysis of Brachyrhaphis by using four molecular markers (3mtDNA, 1nucDNA) to infer relationships among species in this genus. We tested the validity of this genus as a monophyletic group using extensive outgroup sampling based on recent phylogenetic hypotheses of Poeciliidae. We also tested the validity of recently described species of Brachyrhaphis that are part of the B. episcopi complex in Panama. Finally, we examined the impact of historical events on diversification of Brachyrhaphis, and made predictions regarding the role of different ecological environments on evolutionary diversification where known historical events apparently fail to explain speciation. Based on our results, we reject the monophyly of Brachyrhaphis, and question the validity of two recently described species (B. hessfeldi and B. roswithae). Historical biogeography of Brachyrhaphis generally agrees with patterns found in other freshwater taxa in Lower Central America, which show that geological barriers frequently predict speciation. Specifically, we find evidence in support of an 'island' model of Lower Central American formation, which posits that the nascent isthmus was partitioned by several marine connections before linking North and South America. In some cases where historic events (e.g., vicariance) fail to explain allopatric species breaks in Brachyrhaphis, ecological processes (e.g., divergent predation environments) offer additional insight into our understanding of phylogenetic diversification in this group.

Size doesn't matter, sex does: a test for boldness in sister species of Brachyrhaphis fishes.

The effect of divergent natural selection on the evolution of behavioral traits has long been a focus of behavioral ecologists. Predation, due to its ubiquity in nature and strength as a selective agent, has been considered an important environmental driver of behavior. Predation is often confounded with other environmental factors that could also play a role in behavioral evolution. For example, environments that contain predators are often more ecologically complex and "risky" (i.e., exposed and dangerous). Previous work shows that individuals from risky environments are often more bold, active, and explorative than those from low-risk environments. To date, most comparative studies of environmentally driven behavioral divergence are limited to comparisons among populations within species that occur in divergent selective environments but neglect comparisons between species following speciation. This limits our understanding of how behavior evolves post-speciation. The Central American live-bearing fish genus Brachyrhaphis provides an ideal system for examining the relationship between selective environments and behavior, within and between species. Here, we test for differences in boldness between sister species B. roseni and B. terrabensis that occur in streams with and without piscivorous predators, respectively. We found that species do differ in boldness, with species that occur with predators being bolder than those that do not. Within each species, we found that sexes differed in boldness, with males being bolder than females. We also tested for a relationship between size (a surrogate for metabolic rate) and boldness, but found no size effects. Therefore, sex, not size, affects boldness. These results are consistent with the hypothesis that complex and risky environments favor individuals with more bold behavioral traits, but they are not consistent with the hypothesis that size (and therefore metabolic rate) drives divergence in boldness. Finally, our results provide evidence that behavioral trait divergence continues even after speciation is complete.

Effects of predation environment and food availability on somatic growth in the Livebearing Fish Brachyrhaphis rhabdophora (Pisces: Poeciliidae).

Variation in somatic growth rates is of great interest to biologists because of the relationship between growth and other fitness-determining traits, and it results from both genetic and environmentally induced variation (i.e. plasticity). Theoretical predictions suggest that mean somatic growth rates and the shape of the reaction norm for growth can be influenced by variation in predator-induced mortality rates. Few studies have focused on variation in reaction norms for growth in response to resource availability between high-predation and low-predation environments. We used juvenile Brachyrhaphis rhabdophora from high-predation and low-predation environments to test for variation in mean growth rates and for variation in reaction norms for growth at two levels of food availability in a common-environment experiment. To test for variation in growth rates in the field, we compared somatic growth rates in juveniles in high-predation and low-predation environments. In the common-environment experiment, mean growth rates did not differ between fish from differing predation environments, but the interaction between predation environment and food level took the form of a crossing reaction norm for both growth in length and mass. Fish from low-predation environments exhibited no significant difference in growth rate between high and low food treatments. In contrast, fish from high-predation environments exhibited variation in growth rates between high and low food treatments, with higher food availability resulting in higher growth rates. In the field, individuals in the high-predation environment grow at a faster rate than those in low-predation environments at the smallest sizes (comparable to sizes in the common-environment experiment). These data provide no evidence for evolved differences in mean growth rates between predation environments. However, fish from high-predation environments exhibited greater plasticity in growth rates in response to resource availability suggesting that predation environments may exhibit increased variation in food availability for prey fish and consequent selection for plasticity.

Ontogeny and sex alter the effect of predation on body shape in a livebearing fish: sexual dimorphism, parallelism, and costs of reproduction.

Predation can cause morphological divergence among populations, while ontogeny and sex often determine much of morphological diversity among individuals. We used geometric morphometrics to characterize body shape in the livebearing fish Brachyrhaphis rhabdophora to test for interactions between these three major shape-determining factors. We assessed shape variation between juveniles and adults of both sexes, and among adults for populations from high- and low-predation areas. Shape differed significantly between predation regimes for all juveniles regardless of sex. As males grew and matured into adults, ontogenetic shape trajectories were parallel, thus maintaining shape differences in adult males between predation environments. However, shape of adult females between predation environments followed a different pattern. As females grew and matured, ontogenetic shape trajectories converged so that shape differences were less pronounced between mature females in predator and nonpredator environments. Convergence in female body shape may indicate a trade-off between optimal shape for predator evasion versus shape required for the livebearing mode of reproduction.