Raymond D. Semlitsch

Ecosystems are linked through the transfer of materials and energy. Studies examining material fluxes across habitat boundaries frequently quantify unidirectional flows of nutrients and energy. However, material fluxes can be multidirectional, and we lack a conceptual framework to describe how their quantity and stoichiometry influence the net transfer of individual elements between ecosystems. Here we develop a zero net transfer isocline (ZNTI) framework that integrates the relative mass and stoichiometry of fluxes into and out of an ecosystem. We then use case studies with amphibians and salmon to elucidate how life history, ontogenetic shifts in stoichiometry, and trophic interactions shape relative fluxes of nutrients between aquatic and terrestrial ecosystems. Because they increase in both size and Ca content from ova to metamorphs, amphibian life histories strongly bias them towards net Ca export into the terrestrial environment. Because amphibian biomass, C, P, and Ca zero net transfer isoclines do not overlap, there is no value of survivorship where the net flux of biomass, C, P, and Ca are simultaneously balanced between terrestrial and aquatic habitats. The degree of iteroparity and semelparity in salmon strongly affects both the magnitude of net biomass and P flux between riverine and marine environments. While the net direction of biomass flux generally remains strongly biased towards import into the riverine system, net P flux can reach net export into the marine environment because of increasing adult breeding survival leading to reduced mass and %P of what they deposit in rivers (e.g., ova versus whole carcasses). These examples highlight how ontogenetic shifts in body size and stoichiometry result in asymmetric fluxes of elements and biomass that can lead to simultaneous net imports and exports of different elements within the same system. Furthermore, they demonstrate how changes in life-history characteristics and stage-specific survivorship can lead to changes in net elemental transport between ecosystems.

Predator species and body size represent critical factors that have differential effects on prey populations, as well as overall community structure. However, investigations of how morphologically-similar predator species, simultaneous to variation in predator body size, influence lower trophic levels are infrequently performed. We tested whether predator species and body size influenced the functional response curve of three larval ambystomatid salamanders (Ringed Salamander, Ambystoma annulatum Cope, 1886; Spotted Salamander, Ambystoma maculatum (Shaw, 1802); Marbled Salamander, Ambystoma opacum (Gravenhorst, 1807)) while eating congeneric prey. We combined larval salamanders of varying body sizes with up to six prey densities within experimental microcosms. We tested for the shape of the functional response curve and obtained parameter estimates for attack rate and handling time for each predator size – species combination. We found variability among both species and size classes, with a combination of type I and type II functional response curves. Large size classes of predators had higher attack rates than smaller size classes, but equivalently-sized larvae of different species exhibited differences in attack rates and handling time. Our study shows that predation risk varies depending on the size structure and diversity of predators present in a food web, and that grouping predators by either species or size class may reduce the ability to predict changes in community structure resulting from such interactions.

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.

The transfer of energy and nutrients largely depends on the role of animals in the movement of biomass between trophic levels and ecosystems. Despite the historical recognition that amphibians could play an important role in the movement of biomass and nutrients, very few studies have provided reliable estimates of abundance and density of amphibians to reveal their true importance. Here, we provide robust estimates of abundance and density of a dominant species, the Southern Redback Salamander (Plethodon serratus Grobman, 1944), in the oak forest ecosystem of the Ozark Highlands in Missouri. We then use the abundance and density estimates to calculate biomass and nutrient content of salamanders at our study sites in the Ozark forests. Salamanders at the Sinkin Experimental Forest comprise a large amount of protein, energy, and nutrients that greatly exceed estimates derived some 35 years ago in the Hubbard Brook Experimental Forest, New Hampshire. Our estimates (7 300 – 12 900 salamanders·ha⁻¹) are 2–4 times greater than the values reported by Burton and Likens (1975a, Ecology, 56: 1068–1080; 1975b, Copeia, 1975: 541–546). Furthermore, we show that density estimates of other small plethodontid species reported in the literature are nearly an order of magnitude greater than that reported by Burton and Likens. We believe this indicates that previous results have underestimated the importance of salamander biomass, nutrient, and energy flux, and their functional role in regulating invertebrates and carbon retention in forest ecosystems.

Phenology often determines the outcome of interspecific interactions, where early-arriving species often dominate interactions over those arriving later. The effects of phenology on species interactions are especially pronounced in aquatic systems, but the evidence is largely derived from experimental studies. We examined whether differences in breeding phenology between two pond-breeding salamanders (Ambystoma annulatum and A. maculatum) affected metamorph recruitment and demographic traits within natural populations, with the expectation that the fall-breeding A. annulatum would negatively affect the spring-breeding A. maculatum. We monitored populations of each species at five ponds over 4 years using drift fences. Metamorph abundance and survival of A. annulatum were affected by intra- and interspecific processes, whereas metamorph size and date of emigration were primarily influenced by intraspecific effects. Metamorph abundance, snout–vent length, date of emigration and survival for A. maculatum were all predicted by combinations of intra- and interspecific effects, but often showed negative relationships with A. annulatum metamorph traits and abundance. Size and date of metamorphosis were strongly correlated within each species, but in opposite patterns (negative for A. annulatum and positive for A. maculatum), suggesting that the two species use alternative strategies to enhance terrestrial survival and that these factors may influence their interactions. Our results match predictions from experimental studies that suggest recruitment is influenced by intra- and interspecific processes which are determined by phenological differences between species. Incorporating spatiotemporal variability when modeling population dynamics is necessary to understand the importance of phenology in species interactions, especially as shifts in phenology occur under climate change.

Vegetation in and around freshwater ecosystems can affect aquatic organisms through the production of secondary compounds, which are retained in leaves after senescence and are biologically active. Tannins can be toxic to tadpoles, but the plant source of tannins and tannin concentration have been confounded in experimental designs in previous studies. To examine the effects of the concentration and source of tannins (tree species), we examined the effects of 4 factors on tadpole survival, growth, and development: tannin source (red oak [Quercus rubra], white oak [Quercus alba], or sugar maple [Acer saccharum]); tannin concentration (including a control); diet protein level; and tadpole species (American toad [Anaxyrus americanus] and spring peepers [Pseudacris crucifer]). Tannin source and concentration affected spring peeper survival, but American toads had uniformly high survival. Spring peepers had a lower survival rate in high tannin concentrations of oak leachate but a high survival rate in both concentrations of sugar maple leachate. These differences in survival did not correspond with changes in dissolved oxygen, and no effect of dietary protein level on tadpole performance was observed. The presence of plant leachate resulted in increased tadpole growth in both species, but the mechanism for this finding is unclear. The results of the present study show that tannin concentration and source are important factors for tadpole performance, adding further evidence that plant chemistry can affect aquatic organisms. Environ Toxicol Chem 2015;34:120–126. © 2014 SETAC

A better understanding of how individuals respond to variation in habitat quality while moving through heterogeneous habitats is needed to predict ecological phenomena at larger scales, such as local population and metapopulation dynamics. We sought to identify how fine‐scale habitat quality affects the decisions of juvenile pond‐breeding salamanders (Ambystoma maculatumandA. annulatum) to cease dispersive movements away from their natal pond, select a refuge, and settle. Because of the acute susceptibility of juvenile amphibians to evaporative water loss in terrestrial habitats, we predicted that they possess mechanisms for adjusting their behavior in response to variations in fine‐scale habitat quality. We used experimental field enclosures to isolate the effects of habitat quality on settling behavior and employed generalized linear mixed models to examine how manipulations in canopy cover (closed or open) and microhabitat (control, compacted soils, high coarse woody debris, high burrow density), along with environmental variables (rainfall and air temperature), affect the individual's probability of settling. Our results indicated thatA. maculatumandA. annulatumhad a 10% and 30% decreased probability of settling in open‐canopy clearcut habitat, respectively, compared to closed‐canopy forest habitat. In addition,A. annulatumwere 24% less likely to settle in compacted soil treatments. Although the settlement probability ofA. annulatumdid not depend on refuge availability,A. maculatumwere 18% and 25% more likely to settle under conditions of high burrow density and high coarse woody debris, respectively. These findings make a unique contribution to our understanding of amphibian movement ecology by demonstrating how the interplay of external factors and individual behavior produce observed patterns of movement and habitat selection.