Phenotypic shifts caused by predation: selection or life-history shifts?

Predators can impose both selection and life-history shifts in prey populations. Because both processes may affect phenotypic distributions, the estimates of selection differentials may be biased. We carried out two field experiments to disentangle these separate effects. We studied whether dragonfly predation by Aeshna cyanea changes the distributions in body size and lamellae morphology in the damselfly Lestes sponsa. Damselflies have caudal lamellae which are used in escapes by swimming. In a first experiment, we manipulated predator presence (No Aeshna, Encaged Aeshna or Free-ranging Aeshna) and stopped the experiment when all larvae had moulted once. In a second experiment, larvae were confronted with a Free-ranging Aeshna but collected before moulting, and survivors were compared with a control sample taken at the start of the experiment. The presence of Aeshna largely reduced the survival probabilities of the Lestes larvae at a very similar rate in both experiments. Daily survival probabilities did not differ between the No Aeshna and Encaged Aeshna treatments. In the Free-ranging Aeshna treatment of the first experiment, size was reduced compared to the other two treatments, creating a significant apparent selection differential. This was probably mainly due to predator-induced reduced growth because in the second experiment, where growth effects were excluded, size of the survivors did not differ from the control sample. In both experiments there was a significant selection pressure for larger lamellae. Standardized directional selection differentials were similar in both experiments (0.57 and 0.28 phenotypic standard deviation units). No survival selection on lamellae shape was detected. These results are in agreement with previous findings that lamellae size, but not lamellae shape, enhances swimming performance and thereby predator escape in this species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Body size, locomotion, and long bone cross-sectional geometry in indriid primates.

The geometry of the midshaft cross-sections of the femur and humerus of five indriid species was analysed. Internal (marrow cavity) and external diameters were measured on X-rays in the anteroposterior (a-p) and mediolateral (m-l) planes; cross-sectional areas, second moments of area, and section moduli were calculated using formulae for a hollow ellipse. Cortical thickness, robusticity indices (relating external diameters to the length of the bones), and a-p/m-l shape variables were also calculated. Model II regression was supplemented by analyses of correlation between size and shape. Indriids are saltatory, i.e., their locomotion is dominated by the hind limbs. Accordingly, the femur is more rigid than the humerus, and it shows a consistent difference between the a-p and m-l planes in measures related to bending strength. Cortical thickness varies considerably both within and across species. The type specimen of the new species Propithecus tattersalli is virtually indistinguishable from P. verreauxi on the basis of its long bone cross-sectional geometry. Femoral robusticity is uncorrelated with size, but humeral robusticity decreases significantly with increasing size. Femoral shape variables (a-p/m-l) are all negatively correlated with body size, indicating that m-l dimensions of the femur increase at a faster rate than do a-p dimensions. The highly loaded plane of movement seems to be more reinforced in the smaller species. Contrary to static biomechanical scaling predictions of positive allometry, all cross-sectional parameters scale relatively close to isometry. It is concluded that either changes in locomotor performance must compensate for the weight-related increase in forces and moments or that the larger-bodied animals operate appreciably closer to the limits of their safety margins.     

They seem to glide. Are there aerodynamic effects in leaping prosimian primates?

Leaping primates often assume a horizontal position while airborne. When the limbs are spread out in such maneuvers, skin folds between the upper limbs and the trunk are exposed. This has led to the assumption that the animals make use of aerodynamic forces for either gliding, steering, or braking before the landing. In terms of physics, aerodynamic lift or aerodynamic drag can cause the described effects. As coefficients of lift and drag are unknown for flying primates, we have calculated those values that give the animals either a 5% gain or loss in leaping distance. These turn out to be in the range of values for cylinder-shaped "blunt" (unstreamlined) bodies. A significant influence of aerodynamic forces on the flight path can therefore be assumed. The smaller-bodied species (e.g., galagos) are more strongly influenced by their great surface areas. Although frontal areas scale positively allometrically with respect to body mass, air speed gains importance in the larger-bodied species (e.g., sifakas). They cover absolutely greater distances and have the higher takeoff velocities. The actual importance of lift and drag cannot be derived from our theoretical calculations but must be determined experimentally.     

Cannibalism-mediated life history plasticity to combined time and food stress

There is increasing awareness that combinations of biotic and time stress interact in shaping life history plasticity. Despite being widespread and abundant, the role of cannibalism in linking both types of constraints to life history plasticity has been largely neglected. Moreover, no studies disentangled direct (due to the extra meal) and indirect (due to the elimination of the competitor) life history effects of cannibalism, and little is known about their differential dependency on these constraints. We studied effects of cannibalism on the life history of the damselfly Lestes viridis under combinations of time stress (by manipulating the perceived time available in the growth season) and food stress. We reared larvae per two and disentangled direct and indirect effects of cannibalism by preventing cannibalism in half of the cups and by manipulating the per capita food increase after cannibalism. Cannibalism was more frequent under both time stress and food stress and our results show it may help cannibals to compensate for the negative effects of these constraints imposed on life history. Both direct and indirect benefits of cannibalism (increased development and growth rates, larger mass at emergence) were dependent on the timing of cannibalism, being more pronounced or only present when cannibalism occurred early. Moreover, we found that the ecological constraints (time stress and food stress) also differentially shaped some of the direct and indirect effects. Given the differential context-dependency of direct and indirect effects and the fact that direct and indirect life history effects may be both important in shaping life history, disentangling these effects is critical to mechanistically understand under which conditions cannibalism is expected to be adaptive or not.     

Behavioral linkage of pelagic prey and littoral predators: microhabitat selection by Daphnia induced by damselfly larvae

Only recently ecologists started treating the previously separately considered benthic, littoral and pelagic zones of lake ecosystems as closely connected compartments. Here we study a link between organisms belonging to a different compartment – namely the pelagic and the littoral – through behavior in a series of laboratory experiments. Waterfleas of the genus Daphnia are inhabitants of the pelagic zone and suffer a high predation pressure from syntopic vertebrate predators (mainly fish). Presumably to escape this predation, they sometimes migrate in the day to the littoral to seek refuge within macrophytes and return to the pelagic at night. Zygopterans from the genus Ischnura do commonly co-occur in ponds with Daphnia and are known as opportunistic predators of Daphnia . In two initial experiments in microcosms in the lab we showed that Ischnura larvae are littoral predators strongly associated with macrophytes. Although we found that predation rates of individual Ischnura larvae on Daphnia are approximately 1.5 fold lower in macrophytes compared to open water, total predation from Ischnura on Daphnia per unit area is tenfold higher within macrophytes than in open water, making the open water a safer place for Daphnia with regard to Ischnura predation. In a third microcosm experiment we monitored horizontal distribution of Daphnia in the absence, presence and odor only of Ischnura larvae. After 2 hours, on average 10% less Daphnia remained within the vegetation when Ischnura larvae or only their odor were present compared to when Ischnura or their odor were absent. We interpret this as a behavioral anti-predation response of Daphnia to the presence of Ischnura larvae that seems primarily chemically mediated. The observed horizontal migration of the pelagic prey driven by the littoral predator may couple both lake compartments and may interact with the predator–prey relationships within the pelagic.     

A Chaperone in the HSP70 Family Controls Production of Extracellular Fibrils in Myxococcus xanthus

Three independent Tn5-lac insertions in the S1 locus of Myxococcus xanthus inactivate the sglK gene, which is nonessential for growth but required for social motility and multicellular development. The sequence of sglK reveals that it encodes a homologue of the chaperone HSP70 (DnaK). The sglK gene is cotranscribed with the upstream grpS gene, which encodes a GrpE homologue. Unlike sglK, grpS is not required for social motility or development. Wild-type M. xanthus is encased in extracellular polysaccharide filaments associated with the multimeric fibrillin protein. Mutations in sglK inhibit cell cohesion, the binding of Congo red, and the synthesis or secretion of fibrillin, indicating that sglK mutants do not make fibrils. The fibR gene, located immediately upstream of the grpS-sglK operon, encodes a product which is predicted to have a sequence similar to those of the repressors of alginate biosynthesis in Pseudomonas aeruginosa and Pseudomonas putida. Inactivation of fibR leads to the overproduction of fibrillin, suggesting that M. xanthus fibril production and Pseudomonas alginate production are regulated in analogous ways. M. xanthus and Pseudomonas exopolysaccharides may play similar roles in a mechanism of social motility conserved in these gram-negative bacteria.     

Gait mechanics of lemurid primates on terrestrial and arboreal substrates

Aspects of gait mechanics of two lemurid species were explored experimentally. Substrate reaction forces were recorded for three animals each of L. catta and E. fulvus walking and running at voluntary speeds either on a wooden runway with an integrated force platform or on elevated pole supports with a section attached to the force platform. The average height of the back over these substrates and fluctuations in this height were evaluated using video-analysis. Animals preferred walking gaits and lower speeds on the poles, and gallops and higher speeds on the ground. At overlapping speeds, few adjustments to substrate types were identified. Hind limb peak forces are usually lower on the poles than on the ground, and the caudal back is closer to the substrate. This suggests that greater hind limb flexion and reduced limb stiffness occurred on the poles. The support phases for both limbs at higher speeds are slightly elongated on the poles. Forelimb peak forces are not lower, and the trajectory of the caudal back does not follow a smoother path, i.e., not all elements of a compliant gait are present on the simulated arboreal substrates. The horizontal, rigid poles, offered as substitutes for branchlike supports in the natural habitat, may not pose enough of a challenge to require more substantial gait adjustments. Across substrates, forelimb peak forces are generally lower than hind limb peak forces. The interlimb force distribution is similar to that of most other primates with more even limb lengths. Walking gaits present a greater divergence in fore- and hind limb forces than galloping gaits, which are associated with higher forces. The more arboreal E. fulvus has higher forelimb forces than the more terrestrial L. catta, unlike some anthropoid species in which the arborealists have lower forelimb forces than the terrestrialists. As in other primate and nonprimate quadrupeds, the major propulsive thrust comes from the hind limbs in both lemurs. While our data confirm certain aspects of primate gait mechanics (e.g., generally higher hind limb forces), they do not fully support the notion of greater limb compliance. Neither a compliant forelimb on branchlike supports, nor a negative correlation of forelimb force magnitudes with degree of arboreality were observed. Increasing forelimb-to-hind-limb-force-ratios with increasing speed and force magnitudes are also not expected under this paradigm.     

Parallel evolution in ecological and reproductive traits to produce cryptic damselfly species across the holarctic

The damselfly genus Enallagma originated in the Nearctic, and two Nearctic lineages recently underwent radiations partly associated with multiple independent habitat shifts from lakes dominated by fish predators into lakes dominated by dragonfly predators. A previous molecular study of four Palearctic morphospecies and all representative Nearctic species identified the presence of two cryptic species sets, with each set having Palearctic and Nearctic representatives. However, the cryptic species within each set are not sibling species. Here, we present quantitative data on ecologically important larval morphologies and behaviors involved in predator avoidance and on adult male morphological structures involved in mate recognition to quantify the phenotypic relationships among these cryptic species sets. For the adult stage, our data indicate strong parallel evolution of the structures involved in specific mate recognition-the male cerci. For the larval stage, morphometric analyses show that the Palearctic species evolved a nearly identical morphology to the sibling-clade members in the Nearctic that live in waters where dragonflies are the top predators. This implicates the importance of dragonfly predation in the history of the Palearctic clade. Behavioral analyses suggest population differentiation in response to the actual predator environment in the Palearctic clade, consistent with the species differentiation seen in the Nearctic. Our results suggest parallel evolution of adult traits that influence specific mate choice and larval traits that influence ecological performance underlie the striking similarity of Enallagma species across continents. This concurrent parallel evolution in both stages of a complex life cycle, especially when both stages do not share the same selective environment, may be a very unusual mechanism generating cryptic species.     

Accounting for size‐specific predation improves our ability to predict the strength of a trophic cascade

Predation can influence the magnitude of herbivory that grazers exert on primary producers by altering both grazer abundance and their per capita consumption rates via changes in behavior, density‐dependent effects, and size. Therefore, models based solely on changes in abundance may miss key components of grazing pressure. We estimated shifts in grazing pressure associated with changes in the abundance and per capita consumption rates of sea urchins triggered by size‐selective predation by sea otters (Enhydra lutris). Field surveys suggest that sea otters dramatically decreased the abundance and median size of sea urchins. Furthermore, laboratory experiments revealed that kelp consumption by sea urchins varied nonlinearly as a function of urchin size such that consumption rates increased to the 0.56 and 0.68 power of biomass for red and green urchins, respectively. This reveals that shifts in urchin size structure due to size‐selective predation by sea otters alter sea urchin per capita grazing rates. Comparison of two quantitative models estimating total consumptive capacity revealed that a model incorporating shifts in urchin abundance while neglecting urchin size structure overestimated grazing pressure compared to a model that incorporated size. Consequently, incorporating shifts in urchin size better predicted field estimates of kelp abundance compared to equivalent models based on urchin abundance alone. We provide strong evidence that incorporating size‐specific parameters increases our ability to describe and predict trophic interactions.