Implications of species addition and decline for nutrient dynamics in freshwaters.

In terrestrial and aquatic ecosystems, organisms directly affect nutrient storage and cycling by sequestering nutrients via growth and remineralizing nutrients via excretion and egestion. Therefore, species introductions and extirpations can profoundly affect nutrient storage and remineralization rates, and present a challenge for conserving ecosystem function in fresh waters. The literature of consumer-driven nutrient dynamics is growing rapidly, but studies of consumer effects on nutrient storage and remineralization across species and among ecosystems are limited. We compared the effects of 3 grazing taxa, nonnative armored catfish in Mexican streams, native mussels in Oklahoma streams, and native tadpoles in Panamanian streams, on nutrient storage and remineralization. We examined interactions among organismal stoichiometry and biomass, nutrient storage, remineralization rates, and ecosystem size across these 3 groups following species decline (tadpoles and mussels) or introduction (armored catfish) to gain a better understanding of organism-specific effects on nutrient dynamics among freshwater ecosystems. Collectively, our results suggest that the ecosystem-level effect of consumer-driven nutrient dynamics is strongly influenced by environmental variables and is taxon specific. Major changes in biomass of stoichiometrically distinctive organisms can lead to subsequent changes in the flux and storage of elements in an ecosystem, but the overall effect of aquatic animals on nutrient dynamics also is determined by discharge and nutrient-limitation patterns in streams and rivers

Nutrition or Detoxification: Why Bats Visit Mineral Licks of the Amazonian Rainforest

Many animals in the tropics of Africa, Asia and South America regularly visit so-called salt or mineral licks to consume clay or drink clay-saturated water. Whether this behavior is used to supplement diets with locally limited nutrients or to buffer the effects of toxic secondary plant compounds remains unclear. In the Amazonian rainforest, pregnant and lactating bats are frequently observed and captured at mineral licks. We measured the nitrogen isotope ratio in wing tissue of omnivorous short-tailed fruit bats, Carollia perspicillata, and in an obligate fruit-eating bat, Artibeus obscurus, captured at mineral licks and at control sites in the rainforest. Carollia perspicillata with a plant-dominated diet were more often captured at mineral licks than individuals with an insect-dominated diet, although insects were more mineral depleted than fruits. In contrast, nitrogen isotope ratios of A. obscurus did not differ between individuals captured at mineral lick versus control sites. We conclude that pregnant and lactating fruit-eating bats do not visit mineral licks principally for minerals, but instead to buffer the effects of secondary plant compounds that they ingest in large quantities during periods of high energy demand. These findings have potential implications for the role of mineral licks for mammals in general, including humans

Biogeochemical hotspots in Forested Landscapes: The Role of Vernal Pools in Denitrification and Organic Matter

Quantifying spatial and temporal heterogeneity in ecosystem processes presents a challenge for conserving ecosystem function across landscapes. In particular, many ecosystems contain small features that play larger roles in ecosystem processes than their size would indicate; thus, they may represent ‘‘hotspots’’ of activity relative to their surroundings. Biogeochemical hotspots are characterized as small features within a landscape that show comparatively high chemical reaction rates. In northeastern forests in North America, vernal pools are abundant, small features that typically fill in spring with snow melt and precipitation and dry by the end of summer. Ephemeral flooding alters soil moisture and the depth of the soil’s oxic/anoxic boundary, which may affect biogeochemical processes. We studied the effects of vernal pools on leaf-litter decomposition rates, soil enzyme activity, and denitrification in vernal pools to assess whether they function as biogeochemical hotspots. Our results indicate that seasonal inundation enhanced leaf-litter decomposition, denitrification, and enzyme activity in vernal pools relative to adjacent forest sites. Leaves in seasonally flooded areas decomposed faster than leaves in terra firme forest sites. Flooding also influenced the C, N, and P stoichiometry of decomposing leaf litter and explained the variance in microbial extracellular enzyme activity for phosphatase, β-D- glucosidase, and β-N-acetylglucosaminidase. Additionally, denitrification rates were enhanced by seasonal flooding across all of the study pools. Collectively, these data suggest that vernal pool eco- systems may function as hotspots of leaf-litter decomposition and denitrification and play a significant role in decomposition and nutrient dynamics relative to their size

Trophic Trait Evolution Explains Variation in Nutrient Excretion Stoichiometry among Panamanian Armored Catfishes (Loricariidae)

Variation in nutrient excretion rates and stoichiometric ratios (e.g., nitrogen to phosphorus) by consumers can have substantial effects on aquatic ecosystem function. While phylogenetic signals within an assemblage often explain variation in nutrient recycling rates and stoichiometry, the phylogenetically conserved traits that underlie this phenomenon remain unclear. In particular, variation in nutrient excretion stoichiometry across a phylogeny might be driven by phylogenetic patterns in either diet or body stoichiometry. We examined the relative importance of these traits in explaining variation in nutrient recycling rates and stoichiometry in a diverse family of Neotropical-armored catfishes, Loricariidae, in Panamanian streams. We found significant variation in nutrient mineralization traits among species and subfamilies, but variation in nutrient excretion stoichiometry among species was best explained by trophic position rather than body stoichiometry. The variation in trophic position among Panamanian species was consistent with variation in the trophic niche of their genera across South America, suggesting that phylogenetic patterns underpin the evolution of trophic and nutrient excretion traits among these species. Such geographical variation in nutrient mineralization patterns among closely related species may be common, given that trophic variation in fish lineages occurs widely. These results suggest that information on trophic trait evolution within lineages will advance our understanding of the functional contribution of animals to biogeochemical cycling

Behavioral responses of the endemic shrimp Halocardina rubra (Malacostraca:Atyidae) to an introduced fish, Gambusia affinis (Actinopterygii: Poeciliidae) and implications for the trophic structure of Hawaiian anchialine ponds

In the Hawaiian Islands, intentionally introduced exotic fishes have been linked to changes in native biodiversity and community composition. In 1905, the mosquito fish Gambusia affinis was introduced to control mosquitoes. Subsequently, G. affinis spread throughout the Islands and into coastal anchialine ponds. Previous studies suggest that presence of invasive fishes in anchialine ponds may eliminate native species, including the endemic shrimp Halocaridina rubra. We examined effects of G. affinis on H. rubra populations in anchialine ponds on the Kona-Kohala coast of the island of Hawai/i. In the presence of G. affinis, H. rubra exhibited a diel activity pattern that was not seen in fishless ponds. Shrimp in ponds with fish were active only at night. This pattern was evident in anchialine ponds and in laboratory experiments. In laboratory predation experiments, G. affinis preferentially consumed smaller H. rubra, and in the field the H. rubra collected from invaded sites were larger than those from fishless ponds. Analysis of trophic position using stable isotope analyses showed that feeding of H. rubra was not significantly distinct from that of snails, assumed to feed at trophic level 2.0 on epilithic algae, but G. affinis was slightly omnivorous, feeding at tropic level 2.2. The mosquito fish diet was apparently composed primarily of algae when the defensive behavior of H. rubra made them substantially unavailable as prey. The effect of successful establishment of G. affinis on shrimp behavior has the potential to alter abundance of benthic algae and processing and recycling of nutrients in anchialine pond ecosystems

Understanding variation in salamander ionomes: A nutrient balance approach

Ecological stoichiometry uses information on a few key biological elements (C, N, and P) to explain complex ecological patterns. Although factors driving variation in these elements are well-established, expanding stoichiometric principles to explore dynamics of the many other essential elements comprising biological tissues (i.e., the ionome) is needed to determine their metabolic relationships and better understand biological control of elemental flows through ecosystems. 2. In this paper, we report observations of ionomic variation in two species of salamander (Ambystoma opacum and A. talpoideum) across ontogenic stages using specimens from biological collections of two wetlands sampled over a 30-year period. This unique data set allowed us to explore the extent of ionomic variation between species, among ontogenic stages, between sites, and through time. 3. We found species- and to a lesser extent site-specific differences in C, N, and P along with 13 other elements forming salamander ionomes but saw no evidence of temporal changes. Salamander ionomic composition was most strongly related to ontogeny with relatively higher concentrations of many elements in adult males (i.e., Ca, P, S, Mg, Zn, and Cu) compared to metamorphic juveniles, which had greater amounts of C, Fe, and Mn. 4. In addition to patterns of individual elements, covariance among elements was used to construct multi-elemental nutrient balances, which revealed differences in salamander elemental composition between species and sites and changes in elemental proportions across ontogenic development. These multi-elemental balances distinguished among species-site-ontogenic stage groups better than using only C, N, and P. 5. Overall, this study highlights the responsiveness of consumer ionomes to life-history and environmental variation while reflecting underlying relationships among elements tied to biological function. As such, ionomic studies can provide important insights into factors shaping consumer elemental composition and for predicting how these changes might affect higher-order ecological processes

Global Patterns and Controls of Nutrient Immobilization On Decomposing Cellulose In Riverine Ecosystems

Microbes play a critical role in plant litter decomposition and influence the fate of carbon in rivers and riparian zones. When decomposing low-nutrient plant litter, microbes acquire nitrogen (N) and phosphorus (P) from the environment (i.e., nutrient immobilization), and this process is potentially sensitive to nutrient loading and changing climate. Nonetheless, environmental controls on immobilization are poorly understood because rates are also influenced by plant litter chemistry, which is coupled to the same environmental factors. Here we used a standardized, low-nutrient organic matter substrate (cotton strips) to quantify nutrient immobilization at 100 paired stream and riparian sites representing 11 biomes worldwide. Immobilization rates varied by three orders of magnitude, were greater in rivers than riparian zones, and were strongly correlated to decomposition rates. In rivers, P immobilization rates were controlled by surface water phosphate concentrations, but N immobilization rates were not related to inorganic N. The N:P of immobilized nutrients was tightly constrained to a molar ratio of 10:1 despite wide variation in surface water N:P. Immobilization rates were temperature-dependent in riparian zones but not related to temperature in rivers. However, in rivers nutrient supply ultimately controlled whether microbes could achieve the maximum expected decomposition rate at a given temperature

Invasive Fishes Generate Biogeochemical Hotspots in a Nutrient-Limited System

<div><p>Fishes can play important functional roles in the nutrient dynamics of freshwater systems. Aggregating fishes have the potential to generate areas of increased biogeochemical activity, or hotspots, in streams and rivers. Many of the studies documenting the functional role of fishes in nutrient dynamics have focused on native fish species; however, introduced fishes may restructure nutrient storage and cycling freshwater systems as they can attain high population densities in novel environments. The purpose of this study was to examine the impact of a non-native catfish (Loricariidae: <em>Pterygoplichthys</em>) on nitrogen and phosphorus remineralization and estimate whether large aggregations of these fish generate measurable biogeochemical hotspots within nutrient-limited ecosystems. Loricariids formed large aggregations during daylight hours and dispersed throughout the stream during evening hours to graze benthic habitats. Excretion rates of phosphorus were twice as great during nighttime hours when fishes were actively feeding; however, there was no diel pattern in nitrogen excretion rates. Our results indicate that spatially heterogeneous aggregations of loricariids can significantly elevate dissolved nutrient concentrations via excretion relative to ambient nitrogen and phosphorus concentrations during daylight hours, creating biogeochemical hotspots and potentially altering nutrient dynamics in invaded systems.</p> </div

Diel changes in <i>Pterygoplichthys</i> excretion and gut content mass.

<p>Average <i>Pterygoplichthys</i> excretion and gut content mass during daytime (1000–1500 h) and nighttime hours (1900–0400 h). (A) <i>Pterygoplichthys</i> NH₄⁺-N excretion rates; (B) <i>Pterygoplichthys</i> total dissolved phosphorus excretion rates; (C) N:P of <i>Pterygoplichthys</i> excretion; (D) gut content dry mass per wet mass of <i>Pterygoplichthys</i>. Error bars represent ±1 SE.</p

<i>Pterygoplichthys</i> in the Chacamax River (N17°29’047” W91°58’430”).

<p>(A) Daytime aggregation of loricariids. The white line outlines the aggregation boundary. (B) Underwater photo of loricariid aggregation. Individual fish are marked with white numbers (1–35). (C) Loricariids spreading out from aggregation to begin evening feeding. Each dark spot (C) is at least one <i>Pterygoplichthys</i>. A small group of fishes (1–10) have been marked with individual numbers to demonstrate fish abundance. Photo credits: K. A. Capps.</p