Modulation of prey capture kinematics in relation to prey distance helps predict success.

Predators are not perfect, as some of their prey capture attempts result in failure. Successful attempts may be partly due to predators modulating their capture kinematics in relation to variation in the visual cues of the prey to increase the probability of success. In praying mantises, which have been suggested to possess stereoscopic vision, variation in prey distance has been shown to elicit variation in the probability of an attempt. However, it remains to be examined whether variation in prey distance results in mantises modulating their attempt to successfully capture prey. The goals of this study were to examine these relationships using the praying mantis system. Using 11 adult female Sphodromantis lineola, we recorded 192 prey capture attempts at 1000 Hz with two cameras to examine the 3D kinematics of successful and unsuccessful prey capture attempts. Using a combination of principal component analysis (PCA) and logistic regression, our results show that as prey distance increases, mantises adjust through greater and faster expansion of the forelegs and body (PC1), which significantly predicts capture success. However, PC1 only explains 22% of the variation in all prey capture attempts, suggesting that the other components may be related to additional aspects of the prey. Our results suggest that the distances at which mantises prefer to attempt to capture prey may be the result of their greater probability of successfully capturing the prey. These results highlight the range of motions mantises use when attempting to capture prey, suggesting flexibility in their prey capture attempts in relation to prey position.

Thermal tolerance of cyprinids along an urban-rural gradient: Plasticity, repeatability and effects of swimming and temperature shock.

Urbanization changes the thermal profile of streams in much the same way that climate change is predicted to with higher temperatures, more varied flow and rapid temperature pulses with precipitation events. Whether exceptional tolerance to these altered thermal conditions is a pre-requisite for a fish species to inhabit urban streams or if urbanization has changed the thermal physiology of those fish species that persist in urban streams is unknown, but could help predict the outcome of future climate disruption. To test whether residence in urban streams is associated with altered thermal tolerance, we compared thermal tolerance (CTMax) and phenotypic plasticity of thermal tolerance (ΔCTMax/Δ acclimation temperature) in five populations of an urban-tolerant cyprinid, the blacknose dace (Rhinichthys atratulus), from multiple watersheds along an urban/rural gradient. Thermal tolerance of these stream fish was tested while swimming at 10 cm*s-1 but also in static water and after thermal shocks of 4°-6 °C simulating precipitation events. To test whether blacknose dace as a species has unusual thermal tolerance or thermal plasticity, we also compared two blacknose dace populations with two co-resident, co-familiars (creek chub (Semotilus atromaculatus) and rosyside dace (Clinostomus funduloides), that don't persist in urban streams at three different acclimation temperatures. Thermal tolerance of blacknose dace, as measured by a critical thermal maximum test (CTMax), was independent of size and activity level, i.e. individuals had identical thermal tolerance whether swimming or resting and CTMax was significantly repeatable across two levels of activity. Although there was some variance among populations, blacknose dace from streams of varied urbanization generally exhibited comparable thermal tolerances, ability to acclimate to different temperatures and were unaffected by thermal shocks. Rosyside dace had significantly lower thermal tolerance than the other two species but plasticity of thermal tolerance was uniform across the three cyprinid species. Our conclusions are that exceptional thermal tolerance or ability to thermally acclimate are not pre-requisite characters for a given cyprinid species to survive in urban streams, nor has thermal tolerance undergone directional selection in this urban environment.

Evolutionary diversification in the raptorial forelegs of Mantodea: Relations to body size and depth perception.

Limb proportions have evolved among animals to meet functional demands among diverse environments. Studies from terrestrial, vertebrate locomotion have demonstrated that variation in limb proportions have adaptively evolved so animals can perform in a given environment. Most of the research on limb proportion evolution is among vertebrates and terrestrial locomotion, with little information on limb segment evolution in invertebrates or for other functional roles. For example, among invertebrates, multisegmented raptorial forelimbs have evolved multiple times independently to capture prey, but there is little information on the adaptive evolution and diversity of these limbs. Furthermore, as feeding performance is influenced by the sensory system, few studies have examined the coevolution of sensory-motor systems. Using mantises (Mantodea) I examined forelimb diversification among 97 species with a combination of methods, including ternary plots for morphospace visualization, phylogenetically informed allometric relationships, and comparison of evolutionary rates of diversification. Furthermore, using head width as a proxy for depth perception, I examined the correlated evolution of foreleg diversity with depth perception. The results show that among the three segments of the foreleg, the tibia is the smallest, most diverse, and has the highest rate of evolution after body size corrections. Furthermore, while all foreleg segments were related to head width, head width explained the most variation in tibial length compared with other foreleg segments. The results suggest a potential adaptive functional role of tibia length related to the displacement or force produced in this mechanical lever. Furthermore, results from this study support distinct ecomorphs of mantises, as several independent evolutions to grass mimicry evolve similar morphologies. RESEARCH HIGHLIGHT: This study demonstrates interspecific variation among segments of an invertebrate raptorial foreleg. Among Mantodea species the tibia is the most diverse and is related to a proxy for depth perception, while the other segments had strong relationships with body size. This suggests an adaptive, functional role of the tibia during prey capture.

Convergent evolution of locomotor morphology but not performance in gymnotiform swimmers.

Convergent evolution of a novel locomotor strategy implies that a fitness benefit may be associated with the new gait. Opportunities to study this phenomenon are often constrained by a lack of transitional taxa, but teleost fishes offer examples of extant species across such evolutionary shifts in gait. For instance, one species from Osteoglossiformes and the entire order of Gymnotiformes independently evolved a novel gait, gymnotiform locomotion, where thrust is produced by the undulation of an elongate anal fin. Here, we investigate whether this convergence in gait is also associated with similarities in shape, burst swimming abilities, and/or steady-swimming energetics. Specifically, we measured body and fin morphology of fish within Gymnotiformes and Osteoglossiformes, along with closely related Siluriformes and Cypriniformes, to examine the link between gymnotiform locomotion and morphology in a phylogenetic context. Second, we tested the burst swimming capabilities and oxygen consumption during endurance swimming of a subset of the same gymnotiform, osteoglossiform, and cypriniform species, including "transitional" Osteoglossiformes that exhibit intermediate gaits, to determine whether the evolution of this specialized gait is associated with a change in either of these performance metrics. Our results suggest that convergence on the gymnotiform gait is associated with morphological convergence, but does not constrain a fish's maximum sprinting speeds or their energetic demands during steady swimming.

Identifying New Small Proteins in Escherichia coli.

The number of small proteins (SPs) encoded in the Escherichia coli genome is unknown, as current bioinformatics and biochemical techniques make short gene and small protein identification challenging. One method of small protein identification involves adding an epitope tag to the 3' end of a short open reading frame (sORF) on the chromosome, with synthesis confirmed by immunoblot assays. In this study, this strategy was used to identify new E. coli small proteins, tagging 80 sORFs in the E. coli genome, and assayed for protein synthesis. The selected sORFs represent diverse sequence characteristics, including degrees of sORF conservation, predicted transmembrane domains, sORF direction with respect to flanking genes, ribosome binding site (RBS) prediction, and ribosome profiling results. Of 80 sORFs, 36 resulted in encoded synthesized proteins-a 45% success rate. Modeling of detected versus non-detected small proteins analysis showed predictions based on RBS prediction, transcription data, and ribosome profiling had statistically-significant correlation with protein synthesis; however, there was no correlation between current sORF annotation and protein synthesis. These results suggest substantial numbers of small proteins remain undiscovered in E. coli, and existing bioinformatics techniques must continue to improve to facilitate identification.

Variation and repeatability of cutaneous water loss and skin resistance in relation to temperature and diel variation in the lizard Sceloporus consobrinus.

Variation in rates of water loss has been proposed to be an important mechanism in the survival of terrestrial organisms, as high rates of water loss in desiccating environments may lead to hydric stress and death. Vapor density deficit, the driving force for evaporative water loss, increases exponentially as temperature increases. Acute temperature changes may be the result of daily behavioral thermoregulation of ectotherms, which may influence the among individual variation rates of water loss. The goals of this study were to determine (1) how rates of cutaneous water loss (CWL) and skin resistance (Rs) are affected by acute temperature acclimation, (2) how rates of CWL and Rs vary throughout the day allowing behavioral thermoregulation and (3) the repeatability of CWL and Rs within and among sampling periods. We measured CWL and calculated skin resistance (Rs) of 30 male Sceloporus consobrinus lizards across three summers. We measured CWL on the dorsal and ventral surface of each lizard at 23 °C followed by measurements at 35 °C, and three separate times throughout the day. We found a significant increase in Rs and decrease in CWL at increased acclimation temperatures (35 °C), a significant difference in CWL and Rs throughout the day allowing behavioral thermoregulation, and support for the repeatability of CWL and Rs. Our results demonstrate variability in CWL and Rs in relation to temperature acclimation and thermoregulation, but mixed evidence for repeatability across treatments. Our results suggest other factors, such as peripheral blood flow, may be influencing the inter-individual variation in CWL and Rs.

Patterns of variation in feeding strike kinematics of juvenile ghost praying mantis (Phyllocrania paradoxa): are components of the strike stereotypic?

Functional systems, such as feeding mechanics, often involve the evolution of several components of the musculoskeletal system that are moved in coordination to capture prey. Because these systems often involve the quick movement of several structures, some feeding systems have been hypothesized to be stereotypic. While the motor activity patterns are often stereotyped, the subsequent kinematics can be variable, many times in response to variation in prey stimulus (e.g. prey position). Patterns of feeding kinematics have been well studied among vertebrates, with less attention on invertebrate systems. The goal of this study was to examine the amount of stereotypy in the feeding strike kinematics of praying mantises. We filmed eight juvenile ghost praying mantises (Phyllocrania paradoxa) at 1000 Hz across several days within instar 7. We digitized several points that represent the movement of the coxa, trochanter-femur and tibia of the raptorial foreleg to obtain a set of kinematics including angles and angular velocities of the joint, as well as body lunge. Using the coefficient of variation, we found less stereotypy in the approach stage of the strike compared with the sweep. Using Bonferroni-corrected Pearson's correlations of kinematics with prey position, we found few traits related to prey position with the exception of some kinematics of the coxa joint and the amount of lunge used during the strike. Our results suggest that several components of the praying mantis strike are stereotypic, while others exhibit flexibility to ensure successful capture of the prey.

Body ram, not suction, is the primary axis of suction-feeding diversity in spiny-rayed fishes.

Suction-feeding fishes exhibit diverse prey-capture strategies that vary in their relative use of suction and predator approach (ram), which is often referred to as the ram-suction continuum. Previous research has found that ram varies more than suction distance among species, such that ram accounts for most differences in prey-capture behaviors. To determine whether these findings hold at broad evolutionary scales, we collected high-speed videos of 40 species of spiny-rayed fishes (Acanthomorpha) feeding on live prey. For each strike, we calculated the contributions of suction, body ram (swimming) and jaw ram (mouth movement relative to the body) to closing the distance between predator and prey. We confirm that the contribution of suction distance is limited even in this phylogenetically and ecologically broad sample of species, with the extreme suction area of prey-capture space conspicuously unoccupied. Instead of a continuum from suction to ram, we find that variation in body ram is the major factor underlying the diversity of prey-capture strategies among suction-feeding fishes. Independent measurement of the contribution of jaw ram revealed that it is an important component of diversity among spiny-rayed fishes, with a number of ecomorphologies relying heavily on jaw ram, including pivot feeding in syngnathiforms, extreme jaw protruders and benthic sit-and-wait ambush predators. A combination of morphological and behavioral innovations has allowed fish to invade the extreme jaw ram area of prey-capture space. We caution that while two-species comparisons may support a ram-suction trade-off, these patterns do not speak to broader patterns across spiny-rayed fishes.

The evolution of phenotypic plasticity in fish swimming.

Fish have a remarkable amount of variation in their swimming performance, from within species differences to diversity among major taxonomic groups. Fish swimming is a complex, integrative phenotype and has the ability to plastically respond to a myriad of environmental changes. The plasticity of fish swimming has been observed on whole-organismal traits such as burst speed or critical swimming speed, as well as underlying phenotypes such as muscle fiber types, kinematics, cardiovascular system, and neuronal processes. Whether the plastic responses of fish swimming are beneficial seems to depend on the environmental variable that is changing. For example, because of the effects of temperature on biochemical processes, alterations of fish swimming in response to temperature do not seem to be beneficial. In contrast, changes in fish swimming in response to variation in flow may benefit the fish to maintain position in the water column. In this paper, we examine how this plasticity in fish swimming might evolve, focusing on environmental variables that have received the most attention: temperature, habitat, dissolved oxygen, and carbon dioxide variation. Using examples from previous research, we highlight many of the ways fish swimming can plastically respond to environmental variation and discuss potential avenues of future research aimed at understanding how plasticity of fish swimming might evolve. We consider the direct and indirect effects of environmental variation on swimming performance, including changes in swimming kinematics and suborganismal traits thought to predict swimming performance. We also discuss the role of the evolution of plasticity in shaping macroevolutionary patterns of diversity in fish swimming.

Intermediate Kinematics Produce Inferior Feeding Performance in a Classic Case of Natural Hybridization.

Selection on naturally occurring hybrid individuals is a key component of speciation theory, but few studies examine the functional basis of hybrid performance. We examine the functional consequences of hybridization in nature, using the freshwater sunfishes (Centrarchidae), where natural hybrids have been studied for more than a century and a half. We examined bluegill (Lepomis macrochirus), green sunfish (Lepomis cyanellus), and their naturally occurring hybrid, using prey-capture kinematics and morphology to parameterize suction-feeding simulations on divergent parental resources. Hybrid individuals exhibited kinematics intermediate between those of the two parental species. However, performance assays indicated that hybrids display performance most similar to the worse-performing species for a given parental resource. Our results show that intermediate hybrid phenotypes can be impaired by a less-than-intermediate performance and hence suffer a larger loss in fitness than could be inferred from morphology alone.

The evolution of the sexually selected sword in Xiphophorus does not compromise aerobic locomotor performance.

Sexual selection can increase morphological diversity within and among species. Little is known regarding how interspecific variation produced through sexual selection affects other functional systems. Here, we examine how morphological diversity resulting from sexual selection impacts aerobic locomotor performance. Using Xiphophorus (swordtail fish) and their close relatives (N = 19 species), we examined whether the evolution of a longer sexually selected sword affects critical swimming speed. We also examined the effect of other suborganismal, physiological, and morphological traits on critical swimming speed, as well as their relationship with sword length. In correlation analyses, we found no significant relationship between sword length and critical swimming speed. Unexpectedly, we found that critical swimming speed was higher in species with longer swords, after controlling for body size in multiple regression analyses. We also found several suborganismal and morphological predictors of critical swimming speed, as well as a significant negative relationship between sword length and heart and gill mass. Our results suggest that interspecific variation in sword length is not costly for this aspect of swimming performance, but further studies should examine potential costs for other types of locomotion and other components of Darwinian fitness (e.g., survivorship, life span).

Potential enhanced ability of giant squid to detect sperm whales is an exaptation tied to their large body size.

It has been hypothesized that sperm whale predation is the driver of eye size evolution in giant squid. Given that the eyes of giant squid have the size expected for a squid this big, it is likely that any enhanced ability of giant squid to detect whales is an exaptation tied to their body size. Future studies should target the mechanism behind the evolution of large body size, not eye size. Reconstructions of the evolutionary history of selective regime, eye size, optical performance, and body size will improve the understanding of the evolution of large eyes in large ocean animals.

Allometry indicates giant eyes of giant squid are not exceptional.

BACKGROUND: The eyes of giant and colossal squid are among the largest eyes in the history of life. It was recently proposed that sperm whale predation is the main driver of eye size evolution in giant squid, on the basis of an optical model that suggested optimal performance in detecting large luminous visual targets such as whales in the deep sea. However, it is poorly understood how the eye size of giant and colossal squid compares to that of other aquatic organisms when scaling effects are considered. RESULTS: We performed a large-scale comparative study that included 87 squid species and 237 species of acanthomorph fish. While squid have larger eyes than most acanthomorphs, a comparison of relative eye size among squid suggests that giant and colossal squid do not have unusually large eyes. After revising constants used in a previous model we found that large eyes perform equally well in detecting point targets and large luminous targets in the deep sea. CONCLUSIONS: The eyes of giant and colossal squid do not appear exceptionally large when allometric effects are considered. It is probable that the giant eyes of giant squid result from a phylogenetically conserved developmental pattern manifested in very large animals. Whatever the cause of large eyes, they appear to have several advantages for vision in the reduced light of the deep mesopelagic zone.

As the sword grows: individual variation and ontogenetic effects of a sexually selected trait on locomotor performance in Xiphophorus hellerii.

Previous studies aimed at detecting costs of sexually selected traits have yielded mixed results partly because of variable methods. We present a novel approach: a repeated-measures design to examine individual variation in locomotor performance of male Xiphophorus hellerii as the sexually selected sword develops ontogenetically and to determine whether the growth of a sexually selected trait alters consistency of performance. Individual differences in sprint speed, critical swimming speed (stamina), and relative sword length were statistically repeatable over 9 wk. However, using the Akaike Information Criterion corrected for small sample sizes, the best-fit predictive models for swimming performance did not include sword length or relative sword length. Furthermore, in less supported models and within-week comparisons, there was no statistically significant effect of sword length on performance. These results suggest little effect of the sword on locomotor abilities, which is inconsistent with results from some previous experimental manipulations, possibly because compensatory traits develop ontogenetically in parallel with the sword. However, our results are consistent with correlational studies of natural variation that suggest no locomotor cost of the sword. These results do not necessarily imply a complete lack of a cost to the sword but rather lack of a functional cost for swimming performance.

Iterative evolution of increased behavioral variation characterizes the transition to sociality in spiders and proves advantageous.

The evolution of group living is regarded as a major evolutionary transition and is commonly met with correlated shifts in ancillary characters. We tested for associations between social tendency and a myriad of abiotic variables (e.g., temperature and precipitation) and behavioral traits (e.g., boldness, activity level, and aggression) in a clade of spiders that exhibit highly variable social structures (genus Anelosimus). We found that, relative to their subsocial relatives, social species tended to exhibit reduced aggressiveness toward prey, increased fearfulness toward predators, and reduced activity levels, and they tended to occur in warm, wet habitats with low average wind velocities. Within-species variation in aggressiveness and boldness was also positively associated with sociality. We then assessed the functional consequences of within-species trait variation on reconstituted colonies of four test species (Anelosimus eximius, Anelosimus rupununi, Anelosimus guacamayos, and Anelosimus oritoyacu). We used colonies consisting of known ratios of docile versus aggressive individuals and group foraging success as a measure of colony performance. In all four test species, we found that groups composed of a mixture of docile and aggressive individuals outperformed monotypic groups. Mixed groups were more effective at subduing medium and large prey, and mixed groups collectively gained more mass during shared feeding events. Our results suggest that the iterative evolution of depressed aggressiveness and increased within-species behavioral variation in social spiders is advantageous and could be an adaptation to group living that is analogous to the formation of morphological castes within the social insects.

A phylogenetic approach to total evaporative water loss in mammals.

Maintaining appropriate water balance is a constant challenge for terrestrial mammals, and this problem can be exacerbated in desiccating environments. It has been proposed that natural selection has provided desert-dwelling mammals physiological mechanisms to reduce rates of total evaporative water loss. In this study, we evaluated the relationship between total evaporative water loss and body mass in mammals by using a recent phylogenetic hypothesis. We compared total evaporative water loss in 80 species of arid-zone mammals to that in 56 species that inhabit mesic regions, ranging in size from 4 g to 3,500 kg, to test the hypothesis that mammals from arid environments have lower rates of total evaporative water loss than mammals from mesic environments once phylogeny is taken into account. We found that arid species had lower rates of total evaporative water loss than mesic species when using a dichotomous variable to describe habitat (arid or mesic). We also found that total evaporative water loss was negatively correlated with the average maximum and minimum environmental temperature as well as the maximum vapor pressure deficit of the environment. Annual precipitation and the variable Q (a measure of habitat aridity) were positively correlated with total evaporative water loss. These results support the hypothesis that desert-dwelling mammals have lower rates of total evaporative water loss than mesic species after controlling for body mass and evolutionary relatedness regardless of whether categorical or continuous variables are used to describe habitat.

New insights from serranid fishes on the role of trade-offs in suction-feeding diversification.

Suction feeding is central to prey capture in the vast majority of ray-finned fishes and has been well studied from a detailed, mechanistic perspective. Several major trade-offs are thought to have shaped the diversification of suction-feeding morphology and behavior, and have become well established in the literature. We revisited several of these expectations in a study of prey capture morphology and kinematics in 30 species of serranid fishes, a large, ecologically variable group that exhibits diverse combinations of suction and forward locomotion. We find that: (1) diversity among species in the morphological potential to generate suction changes drastically across the range of attack speeds that species use, with all species that use high-speed attacks having low capacity to generate suction, whereas slow-speed attackers exhibit the full range of suction abilities (this pattern indicates a more complex 'ram-suction continuum' than previously recognized); (2) there is no trade-off between the mechanical advantage of the lower jaw opening lever and the speed of jaw depression, revealing that this simple interpretation of lever mechanics fails to predict kinematic diversity; (3) high-speed attackers show increased cranial excursions, potentially to compensate for a decrease in accuracy; (4) the amount of jaw protrusion is positively related to attack speed, but not suction capacity; and (5) a principal component analysis revealed three significant multivariate axes of kinematic variation among species. Two of the three axes were correlated with the morphological potential to generate suction, indicating important but complex relationships between kinematics and suction potential. These results are consistent with other recent studies that show that trade-offs derived from simple biomechanical models may be less of a constraint on the evolutionary diversification of fish feeding systems than previously thought.

Latitudinal and climatic variation in body size and dorsal scale counts in Sceloporus lizards:a phylogenetic perspective.

Squamates often follow an inverse Bergmann's rule, with larger-bodied animals occurring in warmer areas or at lower latitudes. The size of dorsal scales in lizards has also been proposed to vary along climatic gradients, with species in warmer areas exhibiting larger scales, putatively to reduce heat load. We tested for these patterns in the diverse and widespread lizard genus Sceloporus. Among 106 species or populations, body size was associated positively with maximum temperature (consistent with the inverse of Bergmann's rule) and aridity, but did not covary with latitude. Scale size (inferred from the inverse relation with numbers of scales) was positively related to body size. Controlling for body size via multiple regression, scale size was associated negatively with latitude (best predictor), positively with minimum temperature, and negatively with aridity (similar results were obtained using scores from a principal components analysis of latitude and climatic indicators). Thus, lizards with larger scales are not necessarily found in areas with higher temperatures. Univariate analyses indicated phylogenetic signal for body size, scale counts, latitude, and all climate indicators. In all cases, phylogenetic regression models fit the data significantly better than nonphylogenetic models; thus, residuals for log(10) number of dorsal scale rows exhibited phylogenetic signal.

Swimming performance trade-offs across a gradient in community composition in Trinidadian killifish (Rivulus hartii).

The impacts of predation and competition on life history, behavioral, and morphological traits are well established for many organisms, but effects on locomotor performance have received relatively little attention. We examined variation in sprint speed and critical swimming speed (U(crit), a measure of stamina) in the Trinidadian killifish (Rivulus hartii) across a gradient of ecological communities. R. hartii are located in (1) "high-predation" sites with large, piscine piscivores, (2) "Rivulus-guppy" sites with guppies, and (3) "Rivulus-only" sites with only R. hartii. R. hartii suffer higher mortality in high-predation sites. In Rivulus-guppy sites, population densities are reduced and growth rates increased compared with Rivulus-only sites, which likely represent indirect effects of guppy predation on young R. hartii. We show a significant negative relationship, suggesting a trade-off, between sprint speed and endurance; Rivulus from high-predation sites were faster sprinters but had reduced critical swimming speeds. This trade-off was also apparent in correlations of the nine population means. At the individual level, the correlation was weaker and only significantly negative when all nine populations (three from each site) were pooled and values were not corrected for body size. Sex had a significant effect on U(crit), with females having a lower U(crit), but sexes did not differ in sprint speed. Fish from high-predation sites also exhibited increased tail lengths and fineness ratios compared to sites without large predators. The two low-predation sites showed no statistical differences in locomotor performance or morphology.