David R. Chalcraft

PREDATOR IDENTITY AND ECOLOGICAL IMPACTS: FUNCTIONAL REDUNDANCY OR FUNCTIONAL DIVERSITY?

Different species occupy similar trophic positions in natural communities. However, ecologists have often overlooked the consequences of this variation for local communities by assuming that species occupying similar trophic positions are functionally similar. There have been few experimental tests of this important assumption. We tested the assumption of functional similarity by comparing the effect of six different predators (three fish and three salamander species) on an ensemble of larval anuran prey. Our experiment identified substantial variation in the impact of different predators on a variety of responses. Differences among predators in their selection of prey caused the structure of the larval anuran ensemble to vary continuously as opposed to producing discrete alternative states. Predators also differed in their ability to suppress either the total number or biomass of anurans. Thus, performance of larval anurans was dependent upon the identity of the predator. Moreover, the identity of predators judged similar and their degree of similarity depends on the specific response variable. In our study we found no predominance of weak or strong interactors, as predators appear to be evenly distributed along a gradient of interaction strengths. Knowledge of the identity of the species occupying a particular trophic position is crucial to predicting the impact of that trophic position on a community. Ecologists should not simply assume that species are functionally similar. The differing effects of different predators suggest that variation in their distribution across complex landscapes influences prey number, biomass, population dynamics, distribution, and ultimately local and regional species diversity.

SCALE DEPENDENCE IN THE SPECIES-RICHNESS–PRODUCTIVITY RELATIONSHIP: THE ROLE OF SPECIES TURNOVER

Recent research in aquatic systems suggests that productivity-richness re- lationships change with spatial scale and that species turnover (i.e., spatial and temporal variation in species composition) plays an important role in generating this scale depen- dence. The generality of such scale dependence and the effects of variation in temporal scale remain unknown. We examined the extent to which the richness-productivity rela- tionship in terrestrial plant communities depends on spatial or temporal scale and evaluated how spatial and temporal turnover (i.e., species turnover in space and time) generates scale dependence in these relationships using data from two Long-Term Ecological Research (LTER) sites (Jornada and Konza). We found a weak hump-shaped relationship (Jornada) and no relationship (Konza) between richness and productivity at the smallest focal scale (1 m 2 at Jornada and 50 m 2 at Konza) at each site, but strong hump-shaped relationships at the largest focal scale (49 m 2 at Jornada and 200 m 2 at Konza) for each site. Relationships between spatial turnover and productivity at each site mirrored the productivity-richness relationships that emerged at the larger spatial scale (i.e., a significant hump-shaped pattern). In contrast, temporal turnover was unrelated to productivity, and hence increasing temporal scale did not appreciably change the form of the productivity-richness relationship. Our study suggests that the way in which productivity-richness relationships change with spatial or temporal scale depends on the form and strength of the underlying relationship between species turnover and productivity. Moreover, we contend that a dominant effect of increasing productivity is the generation of dissimilarity in species composition among localities that comprise a region, rather than increasing the number of species that occur within local communities. Thus, understanding the mechanisms that cause species turnover to vary with productivity is critical to understanding scale dependence in richness-productivity rela- tionships.

SCALE-DEPENDENT RESPONSES OF PLANT BIODIVERSITY TO NITROGEN ENRICHMENT

Experimental studies demonstrating that nitrogen (N) enrichment reduces plant diversity within individual plots have led to the conclusion that anthropogenic N enrichment is a threat to global biodiversity. These conclusions overlook the influence of spatial scale, however, as N enrichment may alter beta diversity (i.e., how similar plots are in their species composition), which would likely alter the degree to which N-induced changes in diversity within localities translate to changes in diversity at larger scales that are relevant to policy and management. Currently, it is unclear how N enrichment affects biodiversity at scales larger than a small plot. We synthesized data from 18 N-enrichment experiments across North America to examine the effects of N enrichment on plant species diversity at three spatial scales: small (within plots), intermediate (among plots), and large (within and among plots). We found that N enrichment reduced plant diversity within plots by an average of 25% (ranging from a reduction of 61% to an increase of 5%) and frequently enhanced beta diversity. The extent to which N enrichment altered beta diversity, however, varied substantially among sites (from a 22% increase to an 18% reduction) and was contingent on site productivity. Specifically, N enrichment enhanced beta diversity at low-productivity sites but reduced beta diversity at high-productivity sites. N-induced changes in beta diversity generally reduced the extent of species loss at larger scales to an average of 22% (ranging from a reduction of 54% to an increase of 18%). Our results demonstrate that N enrichment often reduces biodiversity at both local and regional scales, but that a focus on the effects of N enrichment on biodiversity at small spatial scales may often overestimate (and sometimes underestimate) declines in regional biodiversity by failing to recognize the effects of N on beta diversity.

EXPERIMENTAL VENUE AND ESTIMATION OF INTERACTION STRENGTH: COMMENT

While experiments are vital for understanding how ecological systems operate, different philosophies exist concerning how experiments should be conducted (e.g., Petranka 1989, Dunham and Beaupre 1998, Resetarits and Fauth 1998, Skelly and Kiesecker 2001; also see the special features in Herpetologica [1989; 45:111– 128] and Ecology [1996; 77:663–705, see Dahler and Strong 1996]). Recently, Skelly (2002) asked how experimental venue (i.e., cattle tanks set up as mesocosms in a field setting vs. screened enclosures placed into natural ponds) influences competitive interactions between two species of larval anurans (Pseudacris crucifer and Rana sylvatica) and how results from the two venues match a standard of realism. He observed that density affected competitive interactions among larval anurans in mesocosms but not in enclosures and concluded that enclosures were more realistic because the observed size of tadpoles measured in the field was more similar to the size of tadpoles predicted by the enclosure experiment than by the mesocosm experiment. Although an empirical examination of venue is valid, we believe that this study has serious flaws and claims differences between venues that erroneously devalue the use of mesocosms. Our goal is to reinterpret the results from Skelly (2002) in light of its design, point out methodological/statistical issues associated with his study, and argue that both venues can make meaningful contributions to the field of ecology if they are designed correctly with regard to the questions being asked and the specific population of interest.

Evaluating the effects of trophic complexity on a keystone predator by disassembling a partial intraguild predation food web

1. Many taxa can be found in food webs that differ in trophic complexity, but it is unclear how trophic complexity affects the performance of particular taxa. In pond food webs, larvae of the salamander Ambystoma opacum occupy the intermediate predator trophic position in a partial intraguild predation (IGP) food web and can function as keystone predators. Larval A. opacum are also found in simpler food webs lacking either top predators or shared prey. 2. We conducted an experiment where a partial IGP food web was simplified, and we measured the growth and survival of larval A. opacum in each set of food webs. Partial IGP food webs that had either a low abundance or high abundance of total prey were also simplified by independently removing top predators and/or shared prey. 3. Removing top predators always increased A. opacum survival, but removal of shared prey had no effect on A. opacum survival, regardless of total prey abundance. 4. Surprisingly, food web simplification had no effect on the growth of A. opacum when present in food webs with a low abundance of prey but had important effects on A. opacum growth in food webs with a high abundance of prey. Simplifying a partial IGP food web with a high abundance of prey reduced A. opacum growth when either top predators or shared prey were removed from the food web and the loss of top predators and shared prey influenced A. opacum growth in a non-additive fashion. 5. The non-additive response in A. opacum growth appears to be the result of supplemental prey availability augmenting the beneficial effects of top predators. Top predators had a beneficial effect on A. opacum populations by reducing the abundance of A. opacum present and thereby reducing the intensity of intraspecific competition. 6. Our study indicates that the effects of food web simplification on the performance of A. opacum are complex and depend on both how a partial IGP food web is simplified and how abundant prey are in the food web. These findings are important because they demonstrate how trophic complexity can create variation in the performance of intermediate predators that play important roles in temporary pond food webs.

Nonconsumptive effects in a multiple predator system reduce the foraging efficiency of a keystone predator

Many studies have demonstrated that the nonconsumptive effect (NCE) of predators on prey traits can alter prey demographics in ways that are just as strong as the consumptive effect (CE) of predators. Less well studied, however, is how the CE and NCE of multiple predator species can interact to influence the combined effect of multiple predators on prey mortality. We examined the extent to which the NCE of one predator altered the CE of another predator on a shared prey and evaluated whether we can better predict the combined impact of multiple predators on prey when accounting for this influence. We conducted a set of experiments with larval dragonflies, adult newts (a known keystone predator), and their tadpole prey. We quantified the CE and NCE of each predator, the extent to which NCEs from one predator alters the CE of the second predator, and the combined effect of both predators on prey mortality. We then compared the combined effect of both predators on prey mortality to four predictive models. Dragonflies caused more tadpoles to hide under leaf litter (a NCE), where newts spend less time foraging, which reduced the foraging success (CE) of newts. Newts altered tadpole behavior but not in a way that altered the foraging success of dragonflies. Our study suggests that we can better predict the combined effect of multiple predators on prey when we incorporate the influence of interactions between the CE and NCE of multiple predators into a predictive model. In our case, the threat of predation to prey by one predator reduced the foraging efficiency of a keystone predator. Consequently, the ability of a predator to fill a keystone role could be compromised by the presence of other predators.

Colonization and saturation of habitats by lizards

We studied the colonization of habitats by four species of iguanid lizard (Urosaurus ornatus, Sceloporus graciosus, S. undulatus, and Uta stansburiana) to determine differences among the species in colonization, differences among habitat types in colonization, and the time trajectory of colonization. We surveyed lizards on eight study plots in three habitats after a density reduction experiment. We discovered no difference among lizard species in their recovery relative to numbers removed. However, there were significant differences in recovery among habitats, suggesting an important role of habitat type in the resilience of the lizards to the perturbation. Colonization by lizards was rapid and most study plots were colonized within 2 to 3 mo of the manipulation. One year after the experiment, all species were represented on all plots. Saturation (ratio of marked lizards to the number removed on each plot) was significantly different between the first and all other post-manipulation census periods. There was no significant difference in saturation among the second (1 yr), third (15 mo) and fourth (27 mo) census periods, suggesting asymptotic saturation within 1 yr of the perturbation. The pattern of colonization by resident lizards was similar to that of all lizards (residents and transients). There was no difference in saturation among species, but there were significant habitat and census period differences in saturation.

Physical and chemical characterization of natural and modified nanoclays and their ecotoxicity on a freshwater algae species (Chlamydomonas reinhardtii): Characterization and toxicity of nanoclays to algal species

Nanoclays represent a class of natural and modified nanomaterials that have received attention from industrial and environmental fields. Studies that assess the physicochemical properties of nanoclays and compare the effects of natural and modified nanoclays are scarce. We assessed the physicochemical characteristics of a natural nanoclay (Na+ montmorillonite) and 2 modified nanoclays (Cloisite® 30B and Novaclay™) in the dry powder state and in solution, and their potential toxic effects on algal population growth (Chlamydomonas reinhardtii). All 3 nanoclays exhibited properties that are thought to cause toxic effects on organisms, but the properties varied among the nanoclays. Cloisite 30B had a low particle stability and a chemical composition that are thought to induce a greater toxic effect on organisms than either Novaclay or natural nanoclay. In contrast, Novaclay and natural nanoclay had a particle shape (nanoplate) in solution that is thought to induce a greater toxic effect on organisms than the type of particle shape (spherical) that Cloisite 30B has in solution. Cloisite 30B suppressed population growth of C. reinhardtii, an effect that increased with dosage. Neither Novaclay nor natural nanoclay affected algal population growth across a broad array of concentrations. The results show that modified nanoclays differ in their impact on algae, and careful thought must be given to their usage because some will have negative consequences if released into aquatic ecosystems. Environ Toxicol Chem 2018;37:2860-2870.

Cooccurrence of prey species alters the impact of predators on prey performance through multiple mechanisms

Abstract When prey are differentially affected by intra and interspecific competition, the cooccurrence of multiple prey species alters the per capita availability of food for a particular prey species which could alter how prey respond to the threat of predation, and hence the overall‐effect of predators. We conducted an experiment to examine the extent to which the nonconsumptive and overall effect of predatory water bugs on snail and tadpole traits (performance and morphology) depended on whether tadpoles and snails cooccurred. Tadpoles and snails differed in their relative susceptibility to intraspecific and interspecific competition, and predators affected both prey species via consumptive and nonconsumptive mechanisms. Furthermore, the overall effect of predators often depended on whether another prey species was present. The reasoning for why the overall effect of predators depended on whether prey species cooccurred, however, differed for each of the response variables. Predators affected snail body growth via nonconsumptive mechanisms, but the change in the overall effect of predators on snail body growth was attributable to how snails responded to competition in the absence of predators, rather than a change in how snails responded to the threat of predation. Predators did not affect tadpole body growth via nonconsumptive mechanisms, but the greater vulnerability of competitively superior prey (snails) to predators increased the strength of consumptive mechanisms (and hence the overall effect) through which predators affected tadpole growth. Predators affected tadpole morphology via nonconsumptive mechanisms, but the greater propensity for predators to kill competitively superior prey (snails) enhanced the ability of tadpoles to alter their morphology in response to the threat of predation by creating an environment where tadpoles had a higher per capita supply of food available to invest in the development of morphological defenses. Our work indicates that the mechanisms through which predators affect prey depends on the other members of the community.