Battle of the Bites: The Effect of Sewage Effluent Exposure on Mosquitofish Biocontrol of Mosquitoes in Residential Louisiana.

Mosquitofish, Gambusia affinis, are eponymous larval mosquito predators. Their ability to colonize and survive in habitats that are uninhabitable by other potential predators allows them to naturally manage larval mosquito populations in most ground pools they are present in. However, effluent from residential onsite wastewater treatment systems (OWTSs) appears to limit the presence of fish predators. This is especially problematic in Louisiana, where regulations allow the discharge of OWTS effluent into open drainage conveyances. To determine the effect of effluent on the capacity of mosquitofish for biocontrol in contaminated areas, we assessed the body condition metrics of populations from two effluent-exposed sites and two sites not exposed to effluent, determined the lethal effect of effluent-contaminated drainage water on fish, and measured the prey consumption rates in the presence of effluent. Female fish collected from effluent-impacted sites had a reduced somatic body condition and most females examined displayed masculinized anal fins resembling the male gonopodium structure. This trait was not seen in fish collected from the control sites and has not yet been documented in association with OWTSs or in the state of Louisiana. Fish from the control sites survived at effluent-contaminated water levels < 70%, and the prey clearance rates increased with dilution. Onsite wastewater treatment system effluent has significant effects on both the short- and long-term persistence of mosquitofish, their body composition, reproductive health, and larval mosquito consumption. These effects likely release mosquito larvae from suppression and may increase the threat of mosquito-transmitted pathogens in effluent-contaminated locations.

Intraspecific variation in feeding and locomotor kinematics during prey capture in redbreast sunfish (Lepomis auritus).

Biomechanics research often revolves around understanding traits impacting suction feeding performance in fishes, using freshwater ray-finned sunfishes (Family Centrarchidae) as models. However, simultaneous feeding and locomotion kinematics during prey capture are not recorded for many species and there is less information on how these kinematics vary within a species and within individuals. To (1) add to existing data on the prey capture kinematics of centrarchids, (2) assess variation in a species both within and across individuals, and (3) compare morphology and prey capture kinematics of well-sampled centrarchids, we filmed five redbreast sunfish (Lepomis auritus) at 500 fps-1 approaching and striking non-evasive prey. Redbreast approach prey at ~30 cm s-1 and use approximately 70% of their maximum gape size. Traits related to feeding are more repeatable than traits related to locomotion. However, the Accuracy Index (AI) was consistent across individuals (AI = 0.76 ± 0.07). Functionally, redbreast sunfish are more similar to bluegill sunfish but morphologically they fall in the intermediate morphospace alongside green sunfish when compared with other centrarchids. These data show that whole organism outcomes (AI) are similar despite variation present both within and across individuals and demonstrate the importance of considering both interspecific and intraspecific differences in the functional diversity of ecologically and evolutionarily important behaviors such as prey capture.

Beyond Suction-Feeding Fishes: Identifying New Approaches to Performance Integration During Prey Capture in Aquatic Vertebrates.

Organisms are composed of hierarchically arranged component parts that must work together to successfully achieve whole organism functions. In addition to integration among individual parts, some ecological demands require functional systems to work together in a type of inter-system performance integration. While performance can be measured by the ability to successfully accomplish ecologically relevant tasks, integration across performance traits can provide a deeper understanding of how these traits allow an organism to survive. The ability to move and the ability to consume food are essential to life, but during prey capture these two functions are typically integrated. Suction-feeding fishes have been used as a model of these interactions, but it is unclear how other ecologically relevant scenarios might reduce or change integration. To stimulate further research into these ideas, we highlight three contexts with the potential to result in changes in integration and underlying performance traits: (1) behavioral flexibility in aquatic feeding modes for capturing alternative prey types, (2) changes in the physical demands imposed by prey capture across environments, and (3) secondary adaptation for suction prey capture behaviors. These examples provide a broad scope of potential drivers of integration that are relevant to selection pressures experienced across vertebrate evolution. To demonstrate how these ideas can be applied and stimulate hypotheses, we provide observations from preliminary analyses of locally adapted populations of Trinidadian guppies (Poecilia reticulata) capturing prey using suction and biting feeding strategies and an Atlantic mudskipper (Periophthalmus barbarus) capturing prey above and below water. We also include a re-analysis of published data from two species of secondarily aquatic cetaceans, beluga whales (Delphinapterus leucas) and Pacific white-sided dolphins (Lagenorhynchus obliquidens), to examine the potential for secondary adaptation to affect integration in suction prey capture behaviors. Each of these examples support the broad importance of integration between locomotor and feeding performance but outline new ways that these relationships can be important when suction demands are reduced or altered. Future work in these areas will yield promising insights into vertebrate evolution and we hope to encourage further discussion on possible avenues of research on functional integration during prey capture.

Integration between swim speed and mouth size evolves repeatedly in Trinidadian guppies and aligns with suction-feeding fishes.

Well-supported correlations between swim speed and mouth size during prey capture suggest the broad existence of an integrated relationship between locomotion and feeding in suction-feeding fishes. However, the influence of specialization on this relationship is unclear. We used divergent populations of Trinidadian guppies (Poecilia reticulata) to test whether integration during suction is generalizable to a non-suction specialist and whether intraspecific specialization of component systems affects their integration. Guppies from replicate high- and low-predation streams were recorded capturing wild-type zooplankton using suction. Alternative general linear models supported a positive correlation between swim speed and mouth size in derived low-predation populations, suggesting that the relationship can be extended in some cases. High-predation populations lack this integration, which may be the result of direct selection or constraints imposed by selection on locomotion. As guppies invade new habitats they may be evolving a new, integrated performance phenotype from a non-integrated ancestor.

Complex Systems Are More than the Sum of Their Parts: Using Integration to Understand Performance, Biomechanics, and Diversity.

Organisms are comprised of many interacting parts, and an increased number or specialization of those parts leads to greater complexity and the necessity for increased integration (the ability of those parts to perform together and maintain a functioning organism). Although this idea is widely recognized among biologists, organisms are more tangibly studied when those parts are considered independently. This reductionist approach has successfully advanced our understanding of organisms' performance. However, performance of one system might (or might not) be dependent on performance of another system to achieve a relevant outcome, and the mechanism of this dependence is poorly understood. We synthesize the concepts of complexity and integration and discuss their application in a biomechanical context. Capture of prey by predatory fishes is used as an example to highlight the application of these ideas. We provide a theoretical framework for future hypotheses of integration and predict an "integration space" for fishes that is then populated with data extracted from the literature. Additionally, using the kinematics of prey-capture in two species of sculpin (Scorpaeniformes: Cottidae), we show that species exhibit multivariate integration in distinct ways, and that these differences add additional insight into ecological divergence that would not be apparent by considering systems independently. Finally, we discuss new insights into organismal performance gained through the study of integration as an emergent property of kinematic systems working together during a common task. Integration is rarely the trait of interest, but we show that future work should adopt a more holistic approach to understand why and how animals perform complex behaviors.

Modelled three-dimensional suction accuracy predicts prey capture success in three species of centrarchid fishes.

Prey capture is critical for survival, and differences in correctly positioning and timing a strike (accuracy) are likely related to variation in capture success. However, an ability to quantify accuracy under natural conditions, particularly for fishes, is lacking. We developed a predictive model of suction hydrodynamics and applied it to natural behaviours using three-dimensional kinematics of three centrarchid fishes capturing evasive and non-evasive prey. A spheroid ingested volume of water (IVW) with dimensions predicted by peak gape and ram speed was verified with known hydrodynamics for two species. Differences in capture success occurred primarily with evasive prey (64-96% success). Micropterus salmoides had the greatest ram and gape when capturing evasive prey, resulting in the largest and most elongate IVW. Accuracy predicted capture success, although other factors may also be important. The lower accuracy previously observed in M. salmoides was not replicated, but this is likely due to more natural conditions in our study. Additionally, we discuss the role of modulation and integrated behaviours in shaping the IVW and determining accuracy. With our model, accuracy is a more accessible performance measure for suction-feeding fishes, which can be used to explore macroevolutionary patterns of prey capture evolution.

The ontogenetic scaling of bite force and head size in loggerhead sea turtles (Caretta caretta): implications for durophagy in neritic, benthic habitats.

Ontogenetic studies of vertebrate feeding performance can help address questions relevant to foraging ecology. Feeding morphology and performance can either limit access to food resources or open up new trophic niches in both aquatic and terrestrial systems. Loggerhead sea turtles are long-lived vertebrates with complex life histories that are marked by an ontogenetic shift from an oceanic habitat to a coastal neritic habitat, and a transition from soft oceanic prey to hard, benthic prey. Although considered durophagous and strong biters, bite performance has not been measured in loggerheads, nor has the ontogeny of bite performance been characterized. In the present study, we collected measurements of bite force in loggerhead turtles from hatchlings to adults. When subadults reach the body size at which the ontogenetic shift occurs, their crushing capability is great enough for them to consume numerous species of hard benthic prey of small sizes. As loggerheads mature and bite performance increases, larger and harder benthic prey become accessible. Loggerhead bite performance eventually surpasses the crushing capability of other durophagous carnivores, thereby potentially reducing competition for hard benthic prey. The increasing bite performance and accompanying changes in morphology of the head and jaws are likely an effective mechanism for resource partitioning and decreasing trophic competition. Simultaneous measurements of body and head size and the use of non-linear reduced major axis regression show that bite force increases with significant positive allometry relative to body size (straight carapace length, straight carapace width and mass) and head size (head width, height and length). Simple correlation showed that all recorded morphometrics were good predictors of measured bite performance, but an AICc-based weighted regression showed that body size (straight carapace width followed by straight carapace length and mass, respectively) were more likely predictors of bite force than head size morphometrics (head width and head length).

Life in the flow lane: differences in pectoral fin morphology suggest transitions in station-holding demand across species of marine sculpin.

Aquatic organisms exposed to high flow regimes typically exhibit adaptations to decrease overall drag and increase friction with the substrate. However, these adaptations have not yet been examined on a structural level. Sculpins (Scorpaeniformes: Cottoidea) have regionalized pectoral fins that are modified for increasing friction with the substrate, and morphological specialization varies across species. We examined body and pectoral fin morphology of 9 species to determine patterns of body and pectoral fin specialization. Intact specimens and pectoral fins were measured, and multivariate techniques determined the differences among species. Cluster analysis identified 4 groups that likely represent differences in station-holding demand, and this was supported by a discriminant function analysis. Primarily, the high-demand group had increased peduncle depth (specialization for acceleration) and larger pectoral fins with less webbed ventral rays (specialization for mechanical gripping) compared to other groups; secondarily, the high-demand group had a greater aspect ratio and a reduced number of pectoral fin rays (specialization for lift generation) than other groups. The function of sculpin pectoral fins likely shifts from primarily gripping where demand is likely low, to an equal dependence on gripping and negative lift generation where demand is likely high. Specialization of the ventral pectoral fin region for gripping likely contributes to the recent diversification of some species into high-demand habitats.

The integration of locomotion and prey capture in divergent cottid fishes: functional disparity despite morphological similarity.

Many mobile animals rely on the integration of locomotion and feeding to capture prey. Fishes commonly swim up to a prey item and utilize a combination of ram and suction feeding for prey capture. Marine cottids represent a diverse and abundant lineage of fishes that exhibit variation in feeding mode that is related to their mouth morphology. However, little is known regarding the integration of the locomotor and feeding systems during prey capture. We quantified the feeding kinematics, feeding performance and integration of locomotion and feeding in two species of divergent cottids: Blepsias cirrhosus (silver-spotted sculpin) and Oligocottus maculosus (tidepool sculpin). Individuals were caught from sympatric habitats near the Bamfield Marine Sciences Centre on Vancouver Island and filmed with a high-speed video camera (500 Hz) while feeding on amphipod prey. Two principal component axes summarize differences in integration and feeding mode despite similarity in attack velocity and feeding morphology (peak gape, peak cranial elevation and peak jaw protrusion). A greater number of correlations between locomotor and feeding variables in B. cirrhosus, compared with O. maculosus, indicate greater integration. We conclude that traditional measures of attack kinematics do not capture functionally and ecologically relevant differences between species. The mechanisms underlying differences in locomotor strategy likely result from unexplored morphological or ecological differences between species. In cottids, integration is apparent in more basal, subtidal species such as B. cirrhosus, and the need for integration may be superceded by demands from the habitat in more derived, shallow-water species such as O. maculosus.