Spencer J. Ingley

Morphological divergence driven by predation environment within and between species of Brachyrhaphis fishes.

Natural selection often results in profound differences in body shape among populations from divergent selective environments. Predation is a well-studied driver of divergence, with predators having a strong effect on the evolution of prey body shape, especially for traits related to escape behavior. Comparative studies, both at the population level and between species, show that the presence or absence of predators can alter prey morphology. Although this pattern is well documented in various species or population pairs, few studies have tested for similar patterns of body shape evolution at multiple stages of divergence within a taxonomic group. Here, we examine morphological divergence associated with predation environment in the livebearing fish genus Brachyrhaphis. We compare differences in body shape between populations of B. rhabdophora from different predation environments to differences in body shape between B. roseni and B. terrabensis (sister species) from predator and predator free habitats, respectively. We found that in each lineage, shape differed between predation environments, consistent with the hypothesis that locomotor function is optimized for either steady swimming (predator free) or escape behavior (predator). Although differences in body shape were greatest between B. roseni and B. terrabensis, we found that much of the total morphological diversification between these species had already been achieved within B. rhabdophora (29% in females and 47% in males). Interestingly, at both levels of divergence we found that early in ontogenetic development, females differed in shape between predation environments; however, as females matured, their body shapes converged on a similar phenotype, likely due to the constraints of pregnancy. Finally, we found that body shape varies with body size in a similar way, regardless of predation environment, in each lineage. Our findings are important because they provide evidence that the same source of selection can drive similar phenotypic divergence independently at multiple divergence levels.

Technical and conceptual considerations for using animated stimuli in studies of animal behavior

Abstract Rapid technical advances in the field of computer animation (CA) and virtual reality (VR) have opened new avenues in animal behavior research. Animated stimuli are powerful tools as they offer standardization, repeatability, and complete control over the stimulus presented, thereby “reducing” and “replacing” the animals used, and “refining” the experimental design in line with the 3Rs. However, appropriate use of these technologies raises conceptual and technical questions. In this review, we offer guidelines for common technical and conceptual considerations related to the use of animated stimuli in animal behavior research. Following the steps required to create an animated stimulus, we discuss (I) the creation, (II) the presentation, and (III) the validation of CAs and VRs. Although our review is geared toward computer-graphically designed stimuli, considerations on presentation and validation also apply to video playbacks. CA and VR allow both new behavioral questions to be addressed and existing questions to be addressed in new ways, thus we expect a rich future for these methods in both ultimate and proximate studies of animal behavior.

Animal personality as a driver of reproductive isolation

Although interest in the ecological and evolutionary implications of animal personality continues to grow, the role that personality plays in speciation has received only modest attention. Here we explore links between personality and speciation, and offer a framework for addressing some of this fields most interesting questions.

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.

Life on the fly: phylogenetics and evolution of the helicopter damselflies (Odonata, Pseudostigmatidae)

Ingley, S.J., Bybee, S.M., Tennessen, K.J., Whiting, M.F. & Branham, M.A. (2012). Life on the fly: phylogenetics and evolution of the helicopter damselflies (Odonata, Pseudostigmatidae). —Zoologica Scripta, 41, 637–650.

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

Natural selections 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.

Repeated geographic divergence in behavior: a case study employing phenotypic trajectory analyses

Environmental effects on behavior have long been a focus of behavioral ecologists. Among the important drivers of behavior is predation environment, which can include the presence/absence of predators, differences in resource availability, and variation in individual density. Environments with predators are often more ecologically complex and “risky” than those without predators. Populations from these environments are sometimes more active and explorative than populations from low-risk, less complex environments. To date, most comparative studies of behavior are limited to within-species comparisons of populations from divergent environments, but neglect comparisons between species following speciation, thus limiting our understanding of post-speciation behavioral evolution. Brachyrhaphis fishes provide an ideal system for studying correlations between divergent environments and behavior within and between species. Here, we test for differences in two behavioral traits—activity and exploration —between sister species Brachyrhaphis roseni and Brachyrhaphis terrabensis that occur in divergent predation environments. Species differed in activity and exploration, with higher activity and exploration levels in populations that co-occur with predators. Furthermore, we found drainage-by-species interactions, indicating that the nature of divergence varied geographically. Using the recently developed phenotypic trajectory analysis (PTA), we quantified this difference and found that, while the geographically isolated populations of sister species tended to evolve in parallel, the magnitude of divergence between species differed between drainages. Our results highlight the utility of PTA for multivariate behavioral data and corroborate past predictions that complex and risky environments are correlated with increased activity and exploration levels and that divergence continues post-speciation.

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.

Reinventing the leaf: Multiple origins of leaf-like wings in katydids (Orthoptera: Tettigoniidae)

Abstract. Insects have developed incredible means to avoid detection by predators. At least five insect orders have species that resemble leaves. Katydids (Orthoptera : Tettigoniidae) are the most diverse and wide-ranging of the leaf-like insects. At least 14 of the 20 extant katydid subfamilies contain species with leaf-like wings. Although it is undisputed that many katydids resemble leaves, methods for delineating the leaf-like from non-leaf-like forms have varied by author and in many cases are not explicitly stated. We provide a simple ratio method that can be used to differentiate the leaf-like and non-leaf-like forms. Geometric morphometrics were used to validate the ratio method. Leaf-like wings have been independently derived in at least 15 katydid lineages. Relative rates of speciation were found to be greater in the non-leaf-like forms, suggesting that leaf-like wings within Tettigoniidae are not a driver of diversification. Likewise, throughout Tettigoniidae, selection seems to be favouring the transition away from leaf-like wings. However, within the large Phaneropterinae subclade, relative speciation and transition rates between the leaf-like and non-leaf-like forms do not differ significantly.

Social context, but not individual personality, alters immigrant viability in a spider with mixed social structure

Immigrant viability is a major determinant of the realized rate of gene flow across populations. For social organisms, the social context in which immigrants disperse across contrasting environments may have important implications for their viability post dispersal. Here, we use social spiders whose individual personalities as well as group personality compositions vary across sites to test whether the strength of selection against immigrants (i.e. mortality rates) differs depending on whether spiders are transplanted (1) as individuals and remain alone, (2) join pre-existing colonies at their new non-native environment, or (3) move with their native group. We also tested for an effect of individual personality on survival. We found that social context, and not individual personality, affects individual survival in foreign environments with contrasting resource levels. Individuals that were transplanted with their native groups suffered higher mortality rates compared to individuals transplanted as singletons, regardless of whether or not they were assimilated into native colonies. Moving as individuals could thus provide an avenue for ongoing gene flow among populations from different resource environments. We found no depressed performance of control colonies that were transplanted across sites with resource levels similar to each colonys site of origin. These results are at odds with the intuition that dispersing as a group should generally enhance the viability of immigrants, at least in social species. We propose that these results could be explained by a mismatch in the ideal group compositions (personality compositions) favoured in different environments, despite a lack of selection on individual personality traits. These results provide a first glimpse into the relative roles of individual personality and social context in mediating gene flow among populations from divergent environments.