Thomas L. Anderson

Intermediate Pond Sizes Contain the Highest Density, Richness, and Diversity of Pond-Breeding Amphibians

We present data on amphibian density, species richness, and diversity from a 7140-ha area consisting of 200 ponds in the Midwestern U.S. that represents most of the possible lentic aquatic breeding habitats common in this region. Our study includes all possible breeding sites with natural and anthropogenic disturbance processes that can be missing from studies where sampling intensity is low, sample area is small, or partial disturbance gradients are sampled. We tested whether pond area was a significant predictor of density, species richness, and diversity of amphibians and if values peaked at intermediate pond areas. We found that in all cases a quadratic model fit our data significantly better than a linear model. Because small ponds have a high probability of pond drying and large ponds have a high probability of fish colonization and accumulation of invertebrate predators, drying and predation may be two mechanisms driving the peak of density and diversity towards intermediate values of pond size. We also found that not all intermediate sized ponds produced many larvae; in fact, some had low amphibian density, richness, and diversity. Further analyses of the subset of ponds represented in the peak of the area distribution showed that fish, hydroperiod, invertebrate density, and canopy are additional factors that drive density, richness and diversity of ponds up or down, when extremely small or large ponds are eliminated. Our results indicate that fishless ponds at intermediate sizes are more diverse, produce more larvae, and have greater potential to recruit juveniles into adult populations of most species sampled. Further, hylid and chorus frogs are found predictably more often in ephemeral ponds whereas bullfrogs, green frogs, and cricket frogs are found most often in permanent ponds with fish. Our data increase understanding of what factors structure and maintain amphibian diversity across large landscapes.

Life history differences influence the impacts of drought on two pond-breeding salamanders

Drought is a strong density-independent environmental filter that contributes to population regulation and other ecological processes. Not all species respond similarly to drought, and the overall impacts can vary depending on life histories. Such differences can necessitate management strategies that incorporate information on individual species to maximize conservation success. We report the effects of a short-term drought on occupancy and reproductive success of two pond-breeding salamanders that differ in breeding phenology (fall vs. spring breeder) across an active military base landscape in Missouri, USA: We surveyed ~200 ponds for the presence of eggs, larvae, and metamorphs from 2011 to 2013. This period coincided with before, during, and after a severe drought that occurred in 2012. The two species showed contrasting responses to drought, where high reproductive failure (34% of ponds) was observed for the spring breeder during a single drought year. Alternatively, the fall breeder only showed a cumulative 8% failure over two years. The number of breeding ponds available for use in the fall decreased during the drought due to pond drying and/or a lack of re-filling. Estimates of occupancy probability declined for the fall-breeding salamander between 2012 and 2013, whereas occupancy probability estimates of the spring breeder increased post-drought. The presence of fish, hydroperiod, the amount of forest cover surrounding ponds, and canopy cover were all found to affect estimates of occupancy probabilities of each species. Pond clustering (distance to nearest pond and the number of ponds within close proximity), hydroperiod, forest cover, and canopy cover influenced both estimates of colonization and extinction probabilities. Our results show life history variation can be important in determining the relative susceptibility of a species to drought conditions, and that sympatric species experiencing the same environmental conditions can respond differently. Consideration of the spatial network and configuration of habitat patches that act as refuges under extreme environmental conditions will improve conservation efforts, such as the placement of permanent ponds for aquatic organisms. A better awareness of species-specific tolerances to environmental filters such as drought can lead to improved management recommendations to conserve and promote habitat for a greater diversity of species across landscapes of spatially connected populations.

Structure and Dynamics of Ringed Salamander (Ambystoma annulatum) Populations in Missouri

Abstract:  We report a detailed account of the structure and dynamics of five populations of Ringed Salamanders (Ambystoma annulatum Cope, 1886) in Missouri, USA, studied over a 4-yr period. The average date of capture for breeding males varied from 30 August to 2 October, and average date of capture for breeding females varied from 9 September to 7 October. The sex ratio was consistently male biased and averaged 2.05 males to each female. The smallest breeding female was 77 mm snout–vent length ([SVL]; mean = 96.3 mm) and the smallest breeding male was 66 mm SVL (mean = 94.2 mm). Eggs and larvae were present in September and October each year, and the larval period varied from 7 to 9 mo. Juveniles metamorphosed in all years in every pond, albeit in low numbers (6–631). The mean SVL of metamorphosing juveniles among all years and ponds was 39.2 mm. Intra-annual body size of emerging juveniles declined over time; juveniles that metamorphosed early were larger in body size than those metamorphosing later in the summer. Production of juveniles per female averaged 0.76 (range, 0.056–2.929), and survival of juveniles from egg to metamorphosis averaged 0.2% (range, 0.01%–0.75%). Both females and males can reach reproductive maturity at 1 yr of age, but most return at 2–3 yr of age. Management of the Ringed Salamander must focus on all life-history stages; protecting the aquatic stage to maximize the number and fitness of metamorphosing juveniles, managing the terrestrial habitat to maximize survival to first reproduction, and monitoring connectivity to promote metapopulation dynamics.

Habitat traits and species interactions differentially affect abundance and body size in pond-breeding amphibians.

In recent studies, habitat traits have emerged as stronger predictors of species occupancy, abundance, richness and diversity than competition. However, in many cases, it remains unclear whether habitat also mediates processes more subtle than competitive exclusion, such as growth, or whether intra- and interspecific interactions among individuals of different species may be better predictors of size. To test whether habitat traits are a stronger predictor of abundance and body size than intra- and interspecific interactions, we measured the density and body size of three species of larval salamanders in 192 ponds across a landscape. We found that the density of larvae was best predicted by models that included habitat features, while models incorporating interactions among individuals of different species best explained the body size of larvae. Additionally, we found a positive relationship between focal species density and congener density, while focal species body size was negatively related to congener density. We posit that salamander larvae may not experience competitive exclusion and thus reduced densities, but instead compensate for increased competition behaviourally (e.g. reduced foraging), resulting in decreased growth. The discrepancy between larval density and body size, a strong predictor of fitness in this system, also highlights a potential shortcoming in using density or abundance as a metric of habitat quality or population health.

High intraguild predator density induces thinning effects on and increases temporal overlap with prey populations

Intraguild (IG) predator density can alter its effects on intraguild prey populations through several mechanisms, including density-dependent processes that affect IG predator traits such as size or growth that enhance or limit its predatory abilities. We examined whether intraspecific density-dependence altered IG predator traits, as well as the subsequent interspecific effects among its intraguild prey within a larval salamander guild. Four densities of ringed salamanders (Ambystoma annulatum), the IG predator, were combined with the presence/absence of spotted salamanders (A. maculatum), the IG prey, within experimental mesocosms. We modeled the effects of A. annulatum density on both conspecific and heterospecific responses that would be indicative of density-dependent competition and predation, respectively. We also modeled the reciprocal interspecific effects of A. maculatum on A. annulatum. We found that increasing intraspecific density negatively affected morphological traits but not survival of A. annulatum. No interspecific effects of A. maculatum on A. annulatum were observed. Alternatively, traits of A. maculatum showed nonlinear relationships with increasing A. annulatum density. Thinning effects of A. annulatum on A. maculatum were observed, as survival was positively and size negatively related for A. maculatum with IG predator density. The temporal overlap of the IG predator and prey also increased nonlinearly with IG predator density, intensifying the potential encounter rate of the two species. Overall, this study shows that density-dependent processes in IG predators can significantly affect traits of both themselves, as well as IG prey, which could ultimately change whether competition or predation occurs between the two groups.

The geography of spatial synchrony

Spatial synchrony, defined as correlated temporal fluctuations among populations, is a fundamental feature of population dynamics, but many aspects of synchrony remain poorly understood. Few studies have examined detailed geographical patterns of synchrony; instead most focus on how synchrony declines with increasing linear distance between locations, making the simplifying assumption that distance decay is isotropic. By synthesising and extending prior work, we show how geography of synchrony, a term which we use to refer to detailed spatial variation in patterns of synchrony, can be leveraged to understand ecological processes including identification of drivers of synchrony, a long-standing challenge. We focus on three main objectives: (1) showing conceptually and theoretically four mechanisms that can generate geographies of synchrony; (2) documenting complex and pronounced geographies of synchrony in two important study systems; and (3) demonstrating a variety of methods capable of revealing the geography of synchrony and, through it, underlying organism ecology. For example, we introduce a new type of network, the synchrony network, the structure of which provides ecological insight. By documenting the importance of geographies of synchrony, advancing conceptual frameworks, and demonstrating powerful methods, we aim to help elevate the geography of synchrony into a mainstream area of study and application.

Non‐additive effects of intra‐ and interspecific competition between two larval salamanders

Assessment of the relative strengths of intra- and interspecific competition has increased in recent years and is critical to understanding the importance of competition. Yet, whether intra- and interspecific competition can have non-additive effects has rarely been tested. The resulting fitness consequences of such non-additive interactions are important to provide the context necessary to advance our understanding of competition theory. We compared the strength of additive and non-additive intra- and interspecific competition by manipulating densities of a pair of larval salamanders (Ambystoma talpoideum and A. maculatum) in experimental mesocosms within a response surface design. Intraspecific density had the strongest effect on the strength of competition for both species, and few observed comparisons indicated interspecific competition was an important factor in predicting body size, growth or larval period length of either species. Non-additive effects of intra- and interspecific competition influenced some response variables, including size and mass at metamorphosis in A. maculatum, but at a reduced strength compared to intraspecific effects alone. Intraspecific competition was thus the dominant biotic interaction, but non-additive effects also impact the outcome of competition in these species, validating the importance of testing for and incorporating non-additive density effects into competition models.

Predation of Eggs and Recently Hatched Larvae of Endemic Ringed Salamanders (Ambystoma annulatum) by Native and Introduced Aquatic Predators

Abstract:  Predation is a key determinant of pond community structure, yet not all predators are equally effective and not all life stages of potential prey are similarly susceptible. Understanding the effects of native and introduced species is essential to informing management strategies, especially for endemic and species of conservation concern. We examined the effects of five common predators (three native: Central Newts [Notophthalmus viridescens louisianensis], aeshnid dragonfly naiads [Aeshnidae], and Southern Leopard Frog tadpoles [Lithobates sphenocephalus]; and two introduced: Fathead Minnows [Pimephales promelas] and Mosquitofish [Gambusia affinis]) on survival of eggs and recently hatched larvae of Ringed Salamanders (Ambystoma annulatum). We also examined the effect of supplemental food or cover availability on survival at each stage. Predators primarily showed a binary response to eggs, consuming all or none of them. Supplemental food did not influence whether eggs or larvae were consumed. Larvae were consumed by all predator species although the effect varied. The presence of cover did not reduce the impacts of the other predators on larval survival. Overall, the two introduced fish species had a greater impact on survival of the early stages of Ringed Salamanders than did the native predators. Further inquiries into the susceptibility of different life stages and survival will improve conservation strategies for rare and endemic species such as Ringed Salamanders.

Top predators and habitat complexity alter an intraguild predation module in pond communities

Predator diversity and habitat complexity frequently influence species interactions at lower trophic levels, yet their joint investigation has been performed infrequently despite the demonstrated importance of each individual factor. We investigated how different top predators and varying habitat complexity influence the function of an intraguild predation module consisting of two larval salamanders, intraguild predator Ambystoma annulatum and intraguild prey A. maculatum. We manipulated predator food webs and habitat complexity in outdoor mesocosms. Top predators significantly influenced body condition and survival of A. annulatum, but habitat complexity had minimal effects on either response. A three-way interaction among the covariates top predator identity, habitat complexity and A. annulatum survival influenced body condition and survival of A. maculatum via a density-mediated indirect effect. Different top predator combinations had variable effects in different habitat complexity treatments on intraguild predator (A. annulatum) survival that subsequently influenced intraguild prey (A. maculatum) body condition and survival. Future work should consider how different top predators influence other food web components, which should vary due to predator attributes and the physical environments in which they co-occur.

Asymmetric effects of intra- and interspecific competition on a pond-breeding salamander

A fundamental goal of evolutionary ecology is to understand how asymmetric competition influences phenotype expression, yet few studies have quantified the relative effects of intra- and interspecific competition on phenotypes. We examined the effects of intra- and interspecific larval competition on both discrete and continuous phenotype expression of the facultatively paedomorphic mole salamander (Ambystoma talpoideum). We manipulated the density and frequency of larval A. talpoideum and a common competitor, the spotted salamander (A. maculatum), in experimental mesocosms within a response surface design. The production of discrete ontogenetic morphs (metamorphs, paedomorphs, and overwintering larvae) was affected more by intraspecific than interspecific competition, but the effect varied between morphs. Paedomorph and metamorph production were more strongly affected by intraspecific than interspecific competition, while the production of overwintering larvae was affected by each type of competition app...