Weston H. Nowlin

COMPARING RESOURCE PULSES IN AQUATIC AND TERRESTRIAL ECOSYSTEMS

Resource pulses affect productivity and dynamics in a diversity of ecosystems, including islands, forests, streams, and lakes. Terrestrial and aquatic systems differ in food web structure and biogeochemistry; thus they may also differ in their responses to resource pulses. However, there has been a limited attempt to compare responses across ecosystem types. Here, we identify similarities and differences in the causes and consequences of resource pulses in terrestrial and aquatic systems. We propose that different patterns of food web and ecosystem structure in terrestrial and aquatic systems lead to different responses to resource pulses. Two predictions emerge from a comparison of resource pulses in the literature: (1) the bottom-up effects of resource pulses should transmit through aquatic food webs faster because of differences in the growth rates, life history, and stoichiometry of organisms in aquatic vs. terrestrial systems, and (2) the impacts of resource pulses should also persist longer in terrestrial systems because of longer generation times, the long-lived nature of many terrestrial resource pulses, and reduced top-down effects of consumers in terrestrial systems compared to aquatic systems. To examine these predictions, we use a case study of a resource pulse that affects both terrestrial and aquatic systems: the synchronous emergence of periodical cicadas (Magicicada spp.) in eastern North American forests. In general, studies that have examined the effects of periodical cicadas on terrestrial and aquatic systems support the prediction that resource pulses transmit more rapidly in aquatic systems; however, support for the prediction that resource pulse effects persist longer in terrestrial systems is equivocal. We conclude that there is a need to elucidate the indirect effects and long-term implications of resource pulses in both terrestrial and aquatic ecosystems.

ALLOCHTHONOUS SUBSIDY OF PERIODICAL CICADAS AFFECTS THE DYNAMICS AND STABILITY OF POND COMMUNITIES

Periodical cicadas emerge from below ground every 13 or 17 years in North American forests, with individual broods representing the synchronous movement of trillions of individuals across geographic regions. Due to predator satiation, most individuals escape predation, die, and become deposited as detritus. Some of this emergent biomass falls into woodland aquatic habitats (small streams and woodland ponds) and serves as a high-quality allochthonous detritus pulse in early summer. We present results of a two-part study in which we (1) quantified deposition of Brood X periodical cicada detritus into woodland ponds and low-order streams in southwestern Ohio, and (2) conducted an outdoor mesocosm experiment in which we examined the effects of deposition of different amounts of cicada detritus on food webs characteristic of forest ponds. In the mesocosm experiment, we manipulated the amount of cicada detritus input to examine if food web dynamics and stability varied with the magnitude of this allochthonous resource subsidy, as predicted by numerous theoretical models. Deposition data indicate that, during years of periodical cicada emergence, cicada carcasses can represent a sizable pulse of allochthonous detritus to forest aquatic ecosystems. In the mesocosm experiment, cicada carcass deposition rapidly affected food webs, leading to substantial increases in nutrients and organism biomass, with the magnitude of increase dependent upon the amount of cicada detritus. Deposition of cicada detritus impacted the stability of organism functional groups and populations by affecting the temporal variability and biomass minima. However, contrary to theory, stability measures were not consistently related to the size of the allochthonous pulse (i.e., the amount of cicada detritus). Our study underscores the need for theory to further explore consequences of pulsed allochthonous subsidies for food web stability.

Effects of Water Level Fluctuation and Short-Term Climate Variation on Thermal and Stratification Regimes of a British Columbia Reservoir and Lake

ABSTRACT Stratification and thermal regimes of a reservoir with fluctuating water levels were compared to a natural lake of similar morphometry and trophic status over a two-year period (2000–2001) in coastal British Columbia, Canada. We compared the timing and duration of stratification, summer heat budgets and heat fluxes in two morphometrically contrasting basins of Sooke Lake Reservoir and Shawnigan Lake (one shallow and one deep basin per water body). In the second year of the study, a 100-year drought allowed us to compare responses of a reservoir and a lake to contrasting years of climatic conditions. Loss of volume from the reservoir during summer and fall caused stratification and thermal regimes to differ from Shawnigan Lake, but the magnitude of these differences was mediated by basin morphometry. Duration of summer stratification, timing of heat content, and the relative importance of seasonal heat fluxes in the shallow basin of Sooke Lake Reservoir were most different from Shawnigan Lake. While there were no major differences between years for Shawnigan Lake, contrasting years in precipitation and hydrology caused Sooke Lake Reservoir stratification and thermal regimes to differ between years. The magnitude of differences between years was mediated by basin size, with the shallower reservoir basin having greater differences between years. Our results indicate that reservoir physical processes are sensitive to short-term changes in hydrology, and that the combined impacts of short-term climate variation and anthropogenic manipulation of hydrology may be greater in shallow reservoir ecosystems.

Trophic ecology of a nonnative population of suckermouth catfish (Hypostomus plecostomus) in a central Texas spring-fed stream

Introduced suckermouth catfishes (Family Loricariidae) are now established in a number of spring-influenced streams in North America but their impacts on native biota are not well documented. Purposes of this study were to quantify gut contents of the loricariid Hypostomus plecostomus from the San Marcos River in central Texas and to evaluate the role of H. plecostomus in the San Marcos River food web using stable isotope analysis. Gut content analysis indicated that H. plecostomus (N = 36) primarily consumed amorphous detritus (87% in biovolume), filamentous red algae (5.4%), and picoplankton (4.1%). Macrophytes, macroinvertebrates, and fish eggs were not found in gut contents. Stable isotopes demonstrate that H. plecostomus occupies a trophic position indicative of an herbivore and likely utilizes detritus of algal origin. The results from our study suggest that large populations of H. plecostomus in the San Marcos River may directly compete with several native herbivorous fishes and may be disrupting trophic flows and nutrient cycling in spring-influenced streams of central and west Texas.

Context-dependent effects of bluegill in experimental mesocosm communities.

Abstract Most knowledge of direct and indirect effects of zooplanktivorous fish has come from studies in which a treatment with a zooplanktivore is compared to a fishless control. However, effects of a zooplanktivore may be different in the presence of other fish species because the other fish have direct and indirect effects that may alter the effects of the zooplanktivore in question. To test this hypothesis, we conducted a tank mesocosm experiment of 2×2 factorial design in which the presence and absence of bluegill (Lepomis macrochirus) were cross-classified with the presence and absence of a fish assemblage composed of western mosquitofish (Gambusia affinis) and common carp (Cyprinus carpio). The presence of bluegill decreased Daphnia, Ceriodaphnia, cyclopoid copepodids, calanoid copepodids, copepod nauplii, amphipods, gastropods, and notonectids. Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, notonectids, Najas, and Chara were decreased and herbivorous rotifers, turbidity, chlorophyll a, total nitrogen and total phosphorus were increased in the presence of the fish assemblage. Significant bluegill×fish assemblage interaction effects were detected for Daphnia, Ceriodaphnia, cyclopoid copepodids, copepod nauplii, gastropods, and notonectids. Analysis of simple effects for these response variables revealed that all significant bluegill effects in the absence of the fish assemblage were not significant in the presence of the fish assemblage. Our results indicate that the effects of bluegill may be context dependent, or dependent upon the presence of other trophically similar fish species.

Gape limitation, prey size refuges and the top-down impacts of piscivorous largemouth bass in shallow pond ecosystems

Top–down control of phytoplankton biomass through piscivorous fish manipulation has been explored in numerous ecological and biomanipulation experiments. Piscivores are gape-limited predators and it is hypothesized that the distribution of gape sizes relative to distribution of body depths of prey fish may restrict piscivore effects cascading to plankton. We examined the top–down effects of piscivorous largemouth bass on nutrients, turbidity, phytoplankton, zooplankton and fish in ponds containing fish assemblages with species representing a range of body sizes and feeding habits (western mosquitofish, bluegill, channel catfish, gizzard shad and common carp). The experimental design consisted of three replicated treatments: fishless ponds (NF), fish community without largemouth bass (FC), and fish community with largemouth bass (FCB). Turbidity, chlorophyll a, cyclopoid copepodid and copepod nauplii densities were significantly greater in FC and FCB ponds than in NF ponds. However, these response variables were not significantly different in FC and FCB ponds. The biomass and density of shallow-bodied western mosquitofish were reduced and bluegill body depths shifted toward larger size classes in the presence of largemouth bass, but the biomass and density of all other fish species and of the total fish community were unaffected by the presence of largemouth bass. Our results show that top–down impacts of largemouth bass in ecosystems containing small- and deep-bodied fish species may be most intense at the top of the food web and alter the size distribution and species composition of the fish community. However, these top–down effects may not cascade to the level of the plankton when large-bodied benthivorous fish species are abundant.

BOTTOM-UP NUTRIENT AND TOP-DOWN FISH IMPACTS ON INSECT-MEDIATED MERCURY FLUX FROM AQUATIC ECOSYSTEMS

Methyl mercury (MeHg) is one of the most hazardous contaminants in the environment, adversely affecting the health of wildlife and humans. Recent studies have demonstrated that aquatic insects biotransport MeHg and other contaminants to terrestrial consumers, but the factors that regulate the flux of MeHg out of aquatic ecosystems via emergent insects have not been studied. The authors used experimental mesocosms to test the hypothesis that insect emergence and the associated flux of MeHg from aquatic to terrestrial ecosystems is affected by both bottom-up nutrient effects and top-down fish consumer effects. In the present study, nutrient addition led to an increase in MeHg flux primarily by enhancing the biomass of emerging insects whose tissues were contaminated with MeHg, whereas fish decreased MeHg flux primarily by reducing the biomass of emerging insects. Furthermore, the authors found that these factors are interdependent such that the effects of nutrients are more pronounced when fish are absent, and the effects of fish are more pronounced when nutrient concentrations are high. The present study is the first to demonstrate that the flux of MeHg from aquatic to terrestrial ecosystems is strongly enhanced by bottom-up nutrient effects and diminished by top-down consumer effects.

Effects of fish on mercury contamination of macroinvertebrate communities of Grassland ponds

Mercury is an environmental contaminant that negatively affects the health of vertebrate consumers such as fish, birds, and mammals. Although aquatic macroinvertebrates are a key link in the trophic transfer of Hg to vertebrate consumers, Hg contamination in macroinvertebrate communities has not been well studied. The purpose of the present study was to examine how Hg in macroinvertebrate communities is affected by the presence of fish. We sampled macroinvertebrates from five ponds with fish and five ponds without fish, at the Lyndon B. Johnson National Grassland in north Texas, USA. Ponds without fish contained a higher biomass of macroinvertebrates and taxa with higher concentrations of Hg, which led to a higher Hg pool in the macroinvertebrate community. A total of 73% of the macroinvertebrate biomass from ponds without fish was composed of taxa with the potential to emerge and transport Hg out of ponds into terrestrial food webs. The results of the present study suggest that small ponds, the numerically dominant aquatic ecosystems in the United States, may be more at risk for containing organisms with elevated Hg concentrations than has been appreciated.

Deposition and decomposition of periodical cicadas (Homoptera: Cicadidae: Magicicada) in woodland aquatic ecosystems

Abstract Many freshwater ecosystems receive allochthonous resource subsidies from adjacent terrestrial environments. In eastern North American forests, geographic broods of periodical cicadas emerge every 13 to 17 y to breed, and local abundances can sometimes be >300 individuals/m2. Most individuals avoid predation, senesce after breeding, and become a resource pulse for forest ecosystems; some cicada carcasses enter freshwater ecosystems where they represent a detrital resource pulse. Here, we present a 2-part study in which we examined the deposition of cicada detritus into woodland ponds and low-order streams in southwestern Ohio during the emergence of Brood X periodical cicadas. We compared the deposition of nutrients associated with periodical cicada detritus and terrestrial leaf litter into small woodland ponds and low-order streams. We used a laboratory experiment to compare patterns of decomposition and nutrient release of adult periodical cicada carcasses and sycamore leaf litter. Input of periodical cicada detritus to woodland streams and ponds was a function of local cicada emergence densities. Organic C loading to woodland aquatic ecosystems from cicada detritus was substantially less than that from terrestrial leaf litter; however, the higher mass-specific N and P content of cicada material made cicada detritus a relatively important nutrient input. N and P deposited in cicada detritus represented 0.2 to 61% of the N and 0.3 to 50% of the P deposited into woodland aquatic ecosystems via terrestrial leaf litter. Decomposition experiments indicated that cicada detritus was of much higher quality than was sycamore leaf litter; female and male cicada carcasses lost mass at significantly faster rates than sycamore leaves (female k = −0.05/d, male k = −0.04/d, sycamore leaf k = −0.002/d). Release rates of C, N, and P from cicada carcasses were 4, 39, and 150× greater, respectively, than release rates from sycamore leaves. Our study indicates that periodical cicada detritus can represent a substantial allochthonous resource pulse to forested aquatic ecosystems and that cicada detritus is of substantially higher quality than is terrestrial leaf litter. These results suggest that deposition and decomposition of periodical cicada detritus can affect the productivity and dynamics of woodland aquatic ecosystems and that the role of animal-derived resource pulses to ecosystems requires further exploration.

Temporal discontinuity of nutrient limitation in plankton communities

Ideas on how various measures of nutrient limitation relate to plankton biomass and species composition are re-examined. While long-term and multi-lake studies typically focus on determining overall biomass, seasonal studies are more focused toward understanding species composition. We use physiological assays to assess short-term nutrient deficiency of nitrogen and phosphorus in two moderately fertile lakes. While biomass in the lakes was considered to ultimately be limited by total phosphorus, nutrient assays were variable in time. Nutrient ratios (TN:TP, PN:PP, PC:PP and PC:PN) did not predict short-term deficiencies, notably that nitrogen deficiency occurred in these phosphorus-limited lakes. In one of our study lakes, there was a relaxation of phosphorus deficiency despite phosphate concentrations occurring below traditional detection limits. Following this period, there was an autumn bloom of Aphanizomenon flos-aquae. This relationship corresponds with other studies that have found A. flos-aquae to be a poor competitor for phosphorus. In contrast, phosphorus deficiency remained high prior to the autumn diatom bloom in our other study lake. Deficiency measures remain an excellent means of assessing physiological status of plankton communities and provide greater insight into species compositional changes, especially when other potential indicators like dissolved nutrient concentrations are inconclusive. Regardless of the nutrient limitation indicator used for a given study, it is critical to consider the appropriate scale of the measure.