Disentangling morphology and genetics in two voles (Microtus pennsylvanicus and M. ochrogaster) in a region of sympatry.

Species in recent, rapid radiations can be difficult to distinguish from one another due to incomplete sorting of traits, insufficient time for novel morphologies to evolve, and elevated rates of hybridization and gene flow. The vole genus Microtus (58 spp.) is one such system where all three factors are likely at play. In the central United States, the prairie vole, Microtus ochrogaster, and the eastern meadow vole, M. pennsylvanicus, occur in sympatry and can be distinguished on the basis of molar cusp patterns but are known to be exceptionally difficult to distinguish using external morphological characters. Using a combination of morphometrics, pelage color analyses, and phylogenetics, we explored which traits are most effective for species identification and whether these same traits can be used to identify the subspecies M. o. ohionensis. While we were able to identify six traits that differed significantly between M. ochrogaster and M. pennsylvanicus, we also found substantial measurement overlap which limits the utility of these traits for species identification. The subspecies M. o. ohionensis was particularly difficult to distinguish from M. p. pennsylvanicus, and we did not find any evidence that this subspecies forms a distinct genetic clade. Furthermore, the full species M. ochrogaster and M. pennsylvanicus did not form reciprocal clades in phylogenetic analyses. We discuss several possible reasons for these patterns, including unrecognized variation in molar cusp patterns and/or localized hybridization. Overall, our results provide useful information that will aid in the identification of these species and subspecies in the future, and provides a case study of how genetics, morphometrics, and fur color analyses can be used to disentangle signatures of evolutionary history and hybridization.

Plant-animal interactions between carnivorous plants, sheet-web spiders, and ground-running spiders as guild predators in a wet meadow community.

Plant-animal interactions are diverse and widespread shaping ecology, evolution, and biodiversity of most ecological communities. Carnivorous plants are unusual in that they can be simultaneously engaged with animals in multiple mutualistic and antagonistic interactions including reversed plant-animal interactions where they are the predator. Competition with animals is a potential antagonistic plant-animal interaction unique to carnivorous plants when they and animal predators consume the same prey.The goal of this field study was to test the hypothesis that under natural conditions, sundews and spiders are predators consuming the same prey thus creating an environment where interkingdom competition can occur.Over 12 months, we collected data on 15 dates in the only protected Highland Rim Wet Meadow Ecosystem in Kentucky where sundews, sheet-web spiders, and ground-running spiders co-exist. One each sampling day, we attempted to locate fifteen sites with: (a) both sheet-web spiders and sundews; (b) sundews only; and (c) where neither occurred. Sticky traps were set at each of these sites to determine prey (springtails) activity-density. Ground-running spiders were collected on sampling days. DNA extraction was performed on all spiders to determine which individuals had eaten springtails and comparing this to the density of sundews where the spiders were captured.Sundews and spiders consumed springtails. Springtail activity-densities were lower, the higher the density of sundews. Both sheet-web and ground-running spiders were found less often where sundew densities were high. Sheet-web size was smaller where sundew densities were high.The results of this study suggest that asymmetrical exploitative competition occurs between sundews and spiders. Sundews appear to have a greater negative impact on spiders, where spiders probably have little impact on sundews. In this example of interkingdom competition where the asymmetry should be most extreme, amensalism where one competitor experiences no cost of interaction may be occurring.

Foraging modality and plasticity in foraging traits determine the strength of competitive interactions among carnivorous plants, spiders and toads.

Foraging modalities (e.g. passive, sit-and-wait, active) and traits are plastic in some species, but the extent to which this plasticity affects interspecific competition remains unclear. Using a long-term laboratory mesocosm experiment, we quantified competition strength and the plasticity of foraging traits in a guild of generalist predators of arthropods with a range of foraging modalities. Each mesocosm contained eight passively foraging pink sundews, and we employed an experimental design where treatments were the presence or absence of a sit-and-wait foraging spider and actively foraging toad crossed with five levels of prey abundance. We hypothesized that actively foraging toads would outcompete the other species at low prey abundance, but that spiders and sundews would exhibit plasticity in foraging traits to compensate for strong competition when prey were limited. Results generally supported our hypotheses. Toads had a greater effect on sundews at low prey abundances, and toad presence caused spiders to locate webs higher above the ground. Additionally, the closer large spider webs were to the ground, the greater the trichome densities produced by sundews. Also, spider webs were larger with than without toads and as sundew numbers increased, and these effects were more prominent as resources became limited. Finally, spiders negatively affected toad growth only at low prey abundance. These findings highlight the long-term importance of foraging modality and plasticity of foraging traits in determining the strength of competition within and across taxonomic kingdoms. Future research should assess whether plasticity in foraging traits helps to maintain coexistence within this guild and whether foraging modality can be used as a trait to reliably predict the strength of competitive interactions.

An insect-feeding guild of carnivorous plants and spiders: does optimal foraging lead to competition or facilitation?

Carnivorous plants and spiders, along with their prey, are main players in an insect-feeding guild found on acidic, poorly drained soils in disturbed habitat. Darwin's notion that these plants must actively attract the insects they capture raises the possibility that spiders could benefit from proximity to prey hotspots created by the plants. Alternatively, carnivorous plants and spiders may deplete prey locally or (through insect redistribution) more widely, reducing each other's gain rates from predation. Here, we formulate and analyze a model of this guild, parameterized for carnivorous sundews and lycosid spiders, under assumptions of random movement by insects and optimal foraging by predators. Optimal foraging here involves gain maximization via trap investment (optimal web sizes and sundew trichome densities) and an ideal free distribution of spiders between areas with and without sundews. We find no facilitation: spiders and sundews engage in intense exploitation competition. Insect attraction by plants modestly increases sundew gain rates but slightly decreases spider gain rates. In the absence of population size structure, optimal spider redistribution between areas with and without sundews yields web sizes that are identical for all spiders, regardless of proximity to sundews. Web-building spiders have higher gain rates than wandering spiders in this system at high insect densities, but wandering spiders have the advantage at low insect densities. Results are complex, indicating that predictions to be tested empirically must be based on careful quantitative assessment.

Evidence for competition between carnivorous plants and spiders.

Several studies have demonstrated that competition between disparate taxa can be important in determining community structure, yet surprisingly, to our knowledge, no quantitative studies have been conducted on competition between carnivorous plants and animals. To examine potential competition between these taxa, we studied dietary and microhabitat overlap between pink sundews (Drosera capillaris) and wolf spiders (Lycosidae) in the field, and conducted a laboratory experiment examining the effects of wolf spiders on sundew fitness. In the field, we found that sundews and spiders had a high dietary overlap with each other and with the available arthropod prey. Associations between sundews and spiders depended on spatial scale: both sundews and spiders were found more frequently in quadrats with more abundant prey, but within quadrats, spiders constructed larger webs and located them further away from sundews as the total sundew trapping area increased, presumably to reduce competition. Spiders also constructed larger webs when fewer prey were available. In the laboratory, our experiment revealed that spiders can significantly reduce sundew fitness. Our findings suggest that members of the plant and animal kingdoms can and do compete.

Fishing spiders, green sunfish, and a stream-dwelling water strider: male-female conflict and prey responses to single versus multiple predator environments.

Many studies have experimentally addressed the effects of a particular predator species on prey behavior. In nature, however, prey frequently face multiple species of predators that often vary in their predatory mode and in their level of predation risk. Relatively few studies have considered prey responses under these complex conditions. In Kentucky, the stream-dwelling water strider (Aquariusremigis) coexists with many potentially dangerous predators, two of which are the green sunfish (Lepomiscyanellus) and the fishing spider (Dolomedesvittatus). Green sunfish occupy stream pools and attack water striders from below. In contrast, fishing spiders hunt along stream shorelines where they perch on overhanging vegetation or rocks and attack water striders near shore. We compared how A. remigis individuals respond to these two very different predators in pools with one or both predators. The presence of sunfish in pools had strong effects on male water strider behavior, including increased use of three types of refuge from sunfish (riffles, climbing out of the water, sitting on the water but at the edges of pools), decreased activity and a decreased number of aggressive males on the water. Spiders also influenced water strider behavior; male water striders avoided spiders by shifting away from the edges of pools. Comparisons of the effects of the two predator species showed that in general, antipredator responses by male water striders were stronger in pools with fish alone than in those with spiders alone. In the presence of both predators, male water strider behavior (microhabitat use and activity) was generally similar to behavior in the presence of fish alone. In contrast, female water striders showed no significant response to the presence of sunfish, and little response to the presence of spiders. This lack of response could be because females spent much of their time in refuges even in the absence of predators (apparently hiding from harassment by males). Both spiders and fish caused decreases in water strider mating activity. The presence of fish reduced both the number of matings per pool (mating frequency), and mean mating durations. Spiders induced a decrease in mean mating duration, but not in mating frequency. The largest reductions in mating activity occurred in pools with both predators present. Pools with either spiders or fish alone suffered 15-20% water strider mortality during our experiment (versus no mortality in predator-free pools). Extant theory suggests that when prey face conflicting microhabitat responses to two predators (as in this study), the predators should have facilitative effects on predation rates (i.e., prey that avoid one predator are often killed by the other and vice versa). Mortality rates in pools with both predators present, however, were not significantly different from that predicted by a null model of multiple predator effects. The lack of predator facilitation can be explained by the compensatory reductions in water strider activity and mating activity in the presence of both predators.

Direct and indirect effects of multiple enemies on water strider mating dynamics.

Previous studies have shown that green sunfish, Lepomis cyanellus, have strong effects on the activity, habitat use, social interactions and mating dynamics of a stream-dwelling water strider, Aquarius remigis (family Gerridae, hence, gerrids). In nature, however, stream pools often contain not just sunfish and water striders, but also smaller fish such as minnows. Here, we used factorial experiments in seminatural streams to document the direct and indirect effects of sunfish and fathead minnows, Pimephales promelas, on water strider survival and behavior. Sunfish, minnows and gerrids all consume surface prey (here, crickets); thus these three species are potential food competitors. Sunfish eat minnows. Accordingly, the presence of sunfish caused minnows to increase their schooling behavior and shift their activity from the surface toward the bottom substrate. The presence of sunfish was also associated with an increase in the number of missing gerrids, whereas minnows caused relatively little gerrid disappearance. Most interestingly, the presence of minnows decreased the effect of sunfish on gerrid disappearance rates; that is, minnows apparently had an indirect positive effect on water strider survival. We suggest that this indirect positive effect reflects the fact that minnows are alternative prey for sunfish. The effects of sunfish and minnows on gerrid mortality explained the influence of these fish on gerrid behavior. Sunfish caused decreases in male gerrid activity, female availability, mating activity, mating frequency and mating duration. Larger males had a mating advantage over smaller males only in pools with sunfish and no minnows. Sunfish also caused a borderline significant decrease in the large female mating advantage. These results were all observed in previous studies and can be viewed as adaptive responses to predation risk. These patterns were not consistent with the expected effects of sunfish as food competitors with water striders. In contrast, minnows had relatively little influence on water strider behavior and the few significant effects were the opposite of those of sunfish. Minnows caused increases in female activity and in mating duration, a decrease in the large male mating advantage and an increase in the large female mating advantage. These patterns fit the view that minnows caused an increase in gerrid hunger, i.e., that minnows acted as food competitors with gerrids. Finally, planned contrasts against controls showed that, in the presence of both sunfish and minnows, water striders showed no significant behavioral responses to fish (i.e., gerrid behavior in pools with sunfish and minnows did not significantly differ from behavior in fishless pools). The most likely mechanism explaining this pattern is a dilution of sunfish predation risk due to the presence of minnows serving as alternative prey for sunfish.