The effects of manipulation of sedimentary iron and organic matter on sediment biogeochemistry and seagrasses in a subtropical carbonate environment

The microbial metabolism of organic matter (OM) in seagrass beds can create sulfidic conditions detrimental to seagrass growth; iron (Fe) potentially has ameliorating effects through titration of the sulfides and the precipitation of iron-sulfide minerals into the sediment. In this study, the biogeochemical effects of Fe availability and its interplay with sulfur and OM on sulfide toxicity, phosphorous (P) availability, seagrass growth and community structure were tested. The availability of Fe and OM was manipulated in a 2 × 2 factorial experiment arranged in a Latin square, with four replicates per treatment. The treatments included the addition of Fe, the addition of OM, the addition of both Fe and OM as well as no addition. The experiment was conducted in an oligotrophic, iron-deficient seagrass bed. Fe had an 84.5% retention efficiency in the sediments with the concentration of Fe increasing in the seagrass leaves over the course of the experiment. Porewater chemistry was significantly altered with a dramatic decrease in sulfide levels in Fe addition plots while sulfide levels increased in the OM addition treatments. Phosphorus increased in seagrass leaves collected in the Fe addition plots. Decreased sulfide stress was evidenced by heavier δ34S in leaves and rhizomes from plots to which Fe was added. The OM addition negatively affected seagrass growth but increased P availability; the reduced sulfide stress in Fe added plots resulted in elevated productivity. Fe availability may be an important determinant of the impact that OM has on seagrass vitality in carbonate sediments vegetated with seagrasses.

Biotic and abiotic determinants of intermediate-consumer trophic diversity in the Florida everglades

Food-web structure can shape population dynamics and ecosystem functioning and stability. We investigated the structure of a food-web fragment consisting of dominant intermediate consumers (fishes and crayfishes) in the Florida Everglades, using stable isotope analysis to quantify trophic diversity along gradients of primary production (periphyton), disturbance (marsh drying) and intermediate-consumer density (a possible indicator of competition). We predicted that trophic diversity would increase with resource availability and decrease after disturbance, and that competition could result in greater trophic diversity by favouring resource partitioning. Total trophic diversity, measured by niche area, decreased with periphyton biomass and an ordination axis representing several bluegreen algae species. Consumers’ basal resource diversity, estimated by δ13C values, was similarly related to algal community structure. The range of trophic levels (δ15N range) increased with time since the most recent drying and reflooding event, but decreased with intermediate-consumer density, and was positively related to the ordination axis reflecting increases in green algae and decreases in filamentous bluegreen algae. Our findings suggest that algal quality, independent of quantity, influences food-web structure and demonstrate an indirect role of nutrient enrichment mediated by its effects on periphyton palatability and biomass. These results reveal potential mechanisms for anthropogenic effects on Everglades communities.

Biogeochemical Processes on Tree Islands in the Greater Everglades: Initiating a New Paradigm

Scientists’ understanding of the role of tree islands in the Everglades has evolved from a plant community of minor biogeochemical importance to a plant community recognized as the driving force for localized phosphorus accumulation within the landscape. Results from this review suggest that tree transpiration, nutrient infiltration from the soil surface, and groundwater flow create a soil zone of confluence where nutrients and salts accumulate under the head of a tree island during dry periods. Results also suggest accumulated salts and nutrients are flushed downstream by regional water flows during wet periods. That trees modulate their environment to create biogeochemical hot spots and strong nutrient gradients is a significant ecological paradigm shift in the understanding of the biogeochemical processes in the Everglades. In terms of island sustainability, this new paradigm suggests the need for distinct dry-wet cycles as well as a hydrologic regime that supports tree survival. Restoration of historic tree islands needs further investigation but the creation of functional tree islands is promising.

Historic primary producer communities linked to water quality and hydrologic changes in the northern Everglades

The northern Everglades Water Conservation Areas have experienced recent ecological shifts in primary producer community structure involving marl periphyton mats and dense Typha-dominated macrophyte stands. Multiple investigations have identified phosphorus (P) as a driver of primary producer community structure, but effects of water impoundment beginning in the 1950s and changes in water hardness [e.g., (CaCO3)] have also been identified as a concern. In an effort to understand pre-1950, primary producer community structure and identify community shifts since 1950, we measured pigment proxies on three sediment cores collected in Water Conservation Area-2A (WCA-2A) along a phosphorus enrichment gradient. Photosynthetic pigments, sediment total phosphorus content (TP), organic matter, total organic carbon and nitrogen were used to infer historic primary producer communities and changes in water quality and hydrology regulating those communities. Excess 210Pb was used to establish historic dates for the sediment cores. Results indicate the northern area of WCA-2A increased marl deposition and increased algal abundance ca. 1920. This increase in (presumably) calcareous periphyton before intensive agriculture and impoundment suggest canal-derived calcium inputs and to some extent early drainage effects played a role in initiating this community shift. The northern area community then shifted to Typha dominance around 1965. The areas to the south in WCA-2A experienced increased marl deposition and algal abundance around or just prior to 1950s impoundment, the precise timing limited by core age resolution. Continued increases in algal abundance were evident after 1950, coinciding with impoundment and deepening of canals draining into WCA-2A, both likely increasing water mineral and nutrient concentrations. The intermediate site developed a Typha-dominated community ca. 1995 while the southern-most core site WCA-2A has yet to develop Typha dominance. Numerous studies link sediment TP >650 mg P/kg to marsh habitat degradation into Typha-dominance. The northern and intermediate cores where Typha is currently support this previous research by showing a distinct shift in the sediment record to Typha dominance corresponding to sediment TP between 600 and 700 mg P/kg. These temporal and spatial differences are consistent with modern evidence showing water-column gradients in mineral inputs (including Ca, carbonates, and phosphorus) altering primary producer community structure in WCA-2A, but also suggest hydroperiod has an effect on the mechanisms regulating periphyton development and Typha dominance.

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2.  We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3.  The best model included bottom-up and top-down effects among trophic groups and supported top-down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom-up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4.  Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top-down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom-up-only model.

Water Quality and Wet Season Diatom Assemblage Characteristics from the Tamiami Trail Pilot Swales Sites (Everglades National Park, Florida, USA)

A pivotal component of hydrological restoration of the Florida Everglades is the improvement of water conveyance to Everglades National Park by the degradation of the current network of canals, roadways and levees. The Tamiami Trail (L29) road/canal complex represents a major barrier to natural water flows into the park and a variety of modification options for flow improvement are currently being explored, including the installation of spreader swales immediately downstream of culverts conveying water under Tamiami Trail from the L29 canal into Everglades National Park. In this study, we evaluated water column chemistry and wet-season diatom community structure to provide baseline information for use in future monitoring activities related to the proposed Tamiami Trail modifications. Water chemistry showed pronounced fluctuations in response to precipitation and anthropogenically mediated hydrological events. Differences in water quality variables among sites were dampened during periods of inundation, and became more pronounced during periods of low canal stage, suggesting the importance of small-scale mechanisms related to isolation of habitat patches. Diatom assemblages were unexpectedly speciose (127 taxa in 40 samples) compared to typical Everglades assemblages, and spatially heterogeneous in sites associated with concentric areas of dense vegetation immediately downstream of culverts. We also observed significant compositional dissimilarities among transects, indicating that culvert pool and north transect assemblages were substantially influenced by propagule input from the canal and areas to the north, while south transect sites were compositionally similar to typical sawgrass prairie diatom communities. Central transect sites were compositionally intermediate to their north and south counterparts. We propose that the position and spatial extent of this “transitional assemblage” is a sensitive indicator of subtle environmental change related to Tamiami Trail modifications.

Dissolved Black Nitrogen (DBN) in Freshwater Environments

Biomass burning results in the formation and accumulation of pyrogenic products such as black carbon (BC) and black nitrogen (BN) in soils. The ubiquitous presence of pyrogenic products in natural dissolved organic matter (DOM) and potential implications in global carbon cycling have recently been reported. However, little is known about the environmental dynamics or the importance in the global N cycle of dissolved BN (DBN; or heteroaromatic N). Here we report the coupling between DBN and dissolved BC (DBC) in ultrafiltered DOM from six headwater streams across a climatic region of North America, suggesting similar combustion sources, and that DOC may play an important role in the translocation of soil BN to the dissolved phase. The export of potentially recalcitrant riverine DBN to the ocean may affect the biogeochemical cycling of N and possibly the microbial community structure in aquatic environments.

A Suite of Prey Traits Determine Predator and Nutrient Enrichment Effects in a Tri-Trophic Food Chain

Predation, predation risk, and resource quality affect suites of prey traits that collectively impact individual fitness, population dynamics, and community structure. However, studies of multi-trophic level effects generally focus on a single prey trait, failing to capture trade-offs among suites of covarying traits that govern population responses and emergent community patterns. We used structural equation models (SEM) to summarize the non-lethal and lethal effects of crayfish, Procambarus fallax, and phosphorus (P) addition, which affected prey food quality (periphyton), on the interactive effects of behavioral, morphological, developmental, and reproductive traits of snails, Planorbella duryi. Univariate and multivariate analyses suggested trade-offs between production (growth, reproduction) and defense (foraging behavior, shell shape) traits of snails in response to non-lethal crayfish and P addition, but few lethal effects. SEM revealed that non-lethal crayfish effects indirectly limited per capita offspring standing stock by increasing refuge use, slowing individual growth, and inducing snails to produce thicker, compressed shells. The negative effects of non-lethal crayfish on snails were strongest with P addition; snails increased allocation to shell defense rather than growth or reproduction. However, compared to ambient conditions, P addition with non-lethal crayfish still yielded greater per capita offspring standing stock by speeding individual snail growth enabling them to produce more offspring that also grew faster. Increased refuge use in response to non-lethal crayfish led to a non-lethal trophic cascade that altered the spatial distribution of periphyton. Independent of crayfish effects, snails stimulated periphyton growth through nutrient regeneration. These findings illustrate the importance of studying suites of traits that reveal costs associated with inducing different traits and how expressing those traits impacts population and community level processes.

Trajectories of Vegetation Response to Water Management in Taylor Slough, Everglades National Park, Florida

Ecosystem management practices that modify the major drivers and stressors of an ecosystem often lead to changes in plant community composition. This paper examines how closely the trajectory of vegetation change in seasonally-flooded wetlands tracks management-induced alterations in hydrology and soil characteristics. We used trajectory analysis, a multivariate method designed to test hypotheses about rates and directions of community change, to examine vegetation shifts in response to changes in water management practices within the Taylor Slough basin of Everglades National Park. We summarized vegetation data by non-metric multidimensional scaling ordination, and examined the time trajectory of each site along environmental vectors representing hydrology and soil phosphorus gradients. In the Taylor Slough basin, vegetation change trajectories closely followed the hydrologic changes caused by the operation of water pumps and detention ponds adjacent to the canals. We also observed a shift in vegetation composition along a vector of increasing soil phosphorus, which suggests the need for implementing measures to avoid P-enrichment in southern Everglades marl prairies. This study indicates that shifts in vegetation composition in response to changes in hydrologic conditions and associated parameters may be detected through trajectory analysis, thereby providing feedback for adaptive management of wetland ecosystems.

A Geospatial Database of Tree Islands within the Mustang Corner Fire Incident of 2008

Fire, which affects community structure and composition at all trophic levels, is an integral component of the Everglades ecosystem (Wade et al. 1980; Lockwood et al. 2003). Without fire, the Everglades as we know it today would be a much different place. This is particularly true for the short-hydroperiod marl prairies that predominate on the eastern and western flanks of Shark River Slough, Everglades National Park (Figure 1). In general, fire in a tropical or sub-tropical grassland community favors the dominance of C4 grasses over C3 species (Roscoe et al. 2000; Briggs et al. 2005). Within this pyrogenic graminoid community also, periodic natural fires, together with suitable hydrologic regime, maintain and advance the dominance of C4 vs C3 graminoids (Sah et al. 2008), and suppress the encroachment of woody stems (Hanan et al. 2009; Hanan et al. unpublished manuscript) originating from the tree islands that, in places, dominate the landscape within this community. However, fires, under drought conditions and elevated fuel loads, can spread quickly throughout the landscape, oxidizing organic soils, both in the prairie and in the tree islands, and, in the process, lead to shifts in vegetation composition. This is particularly true when a fire immediately precedes a flood event (Herndon et al. 1991; Lodge 2005; Sah et al. 2010), or if so much soil is consumed during the fire that the hydrologic regime is permanently altered as a result of a decrease in elevation (Zaffke 1983).

Diversity Patterns of Amphibians in Lowland Amazonian Forests in Southeastern Peru

Biological diversity is threatened worldwide and it is a priority to generate more information that can be used both for understanding ecological processes and determining conservation strategies. For my dissertation, I focused on amphibian diversity patterns in lowland rainforests of southwestern Amazonia to evaluate the importance of habitat heterogeneity in the region. My main purpose was to test the hypothesis that amphibian communities in different forest types differ in species richness, composition, and abundance. I used standardized visual encounter surveys to quantify the species composition and abundance of amphibians at four sites, each containing four forest types (floodplain, terra firme, bamboo, and palm swamp). I used leaf-litter plots to evaluate the effect of soil and leaf-litter characteristics on species richness and abundance of leaf-litter frogs. I intensively sampled at one site and then sampled three other sites (distance among sites varied 3.5-105 km) to evaluate whether the patterns observed at one site were similar elsewhere. I also updated the information on threatened and potentially threatened amphibians in Peru and my study region. I found that no species appears to have experienced population declines in southeastern Peru, suggesting that the region still contains the original species pool. My results support the hypothesis that amphibian communities differ across forest types and that patterns observed at the local scale (one site) are similar at the regional scale (four sites). My data also indicate that there is no correlation between species composition and geographic distance among sites. Instead, an important proportion of the gamma diversity is represented by habitat-related beta diversity. My leaf-litter plot data showed that part of the variation in the leaf-litter community structure is explained by soil and litter characteristics. I found that soil total phosphorus and, to a lesser extent, humidity, leaf-litter mass, and pH is linked to species presence/absence and abundance. My study provides the first standardized, quantitative comparison of amphibian community structure across four major forest types in southwestern Amazonia and highlights the fact that forest types are complementary and necessary for maintaining high species richness in the region.

Macroinvertebrate abundance in montane tropical streams of varying land-use, Ecuador

Despite the importance of tropical montane cloud forest streams, studies investigating aquatic communities in these regions are rare and knowledge on the driving factors of community structure is missing. The objectives of this study therefore were to understand how land-use influences habitat structure and macroinvertebrate communities in cloud forest streams of southern Ecuador. We evaluated these relationships in headwater streams with variable land cover, using multivariate statistics to identify relationships between key habitat variables and assemblage structure, and to resolve differences in composition among sites. Results show that shading intensity, substrate type and pH were the environmental parameters most closely related to variation in community composition observed among sites. In addition, macroinvertebrate density and partly diversity was lower in forested sites, possibly because the pH in forested streams lowered to almost 5 during spates. Standard bioindicator metrics were unable to detect the changes in assemblage structure between disturbed and forested streams. In general, our results indicate that tropical montane headwater streams are complex and heterogeneous ecosystems with low invertebrate densities. We also found that some amount of disturbance, i.e. patchy deforestation, can lead at least initially to an increase in macroinvertebrate taxa richness of these streams.

A 34-month sediment trap experiment at the Cape Verde Ocean Observatory (CVOO, December 2009 - October 2012)

Anticyclonic mesoscale eddies (ACME) have been proposed as a mechanism by which new nutrients are episodically delivered into the euphotic zone, thereby enhancing new production as well as shifting phytoplankton community structure. In this paper, we report on a 34-month sediment trap experiment at the Cape Verde Ocean Observatory (CVOO; ca. 18°N, 24°E; December 2009-October 2012), occasionally influenced by ACME passages. The typically oligotrophic, weakly seasonal particle flux pattern at the CVOO is strongly modified by the appearance of a highly productive and low oxygen ACME. Out of four recorded diatom flux maxima at CVOO, three were associated with the passage of ACMEs. The recorded diatom maxima events support the view that local ACME dynamics promotes upward nutrient supply into the euphotic zone leading to a rapid response of diatoms. This response is clearly reflected by the flux seasonality: between 40% and 60% of the total annual diatom flux at the CVOO site was intercepted in a relatively short time interval (<60 days). A highly diverse diatom community characterized the diatom fluxes throughout. Along with the ACME passages, small species of the genus Nitzschia, and Thalassionema nitzschioides var. parva dominated and delivered a major portion of the opal and organic carbon into deeper waters at site CVOO. Several pelagic, warm-water background species became dominant during intervals with low nutrient availability in the euphotic zone. Results of our interannual time-series suggest that ACMEs impact on total diatom production and the species-specific composition of the assemblage north of the Cave Verde Islands, and can strengthen the biological pump in open-ocean, oligotrophic subtropical regions of the world ocean. Our observations are useful for testing biogeochemical ocean models and will also help in improving the knowledge of processes and mechanisms behind interannual time-series of bulk components and microorganisms in pelagic and hemipelagic ocean areas.

Impacts of newly established non-indigenous green crab (Carcinus maenas) on native fauna in Placentia Bay, Newfoundland

The recent invasion of the European green crab (Carcinus maenas) populations in Placentia Bay, Newfoundland and Labrador (NL) raises great concern about potential impacts on local fisheries and native biodiversity. Green crab are highly adaptable and in both native and invaded areas, green crab are well established predators that can outcompete other similarly sized decapods. The main objectives of this thesis were to: 1) identify the native species that green crab compete with for resources; 2) determine the depths and substrate types in which these interactions likely occur; 3) assess the indirect effects of green crab on native crustaceans and their changes in behavior; 4) assess the impacts of green crab on benthic community structure; 5) compare the NL population with other Atlantic Canadian populations in terms of competitive abilities; and 6) compare morphological features of the NL population with other Atlantic Canadian populations. I found that green crab overlap in space and diet with both rock crab (Cancer irroratus) and American lobster (Homarus americanus), potentially leading to a shift in habitat. Laboratory studies on naïve juvenile lobster also suggested shifts in behavior related to green crab, in that lobster decreased foraging activity and increased shelter use in the presence of green crab. Benthic community analyses showed fewer species in mud, sand, and eelgrass sites heavily populated by green crab compared to sites without green crab, although results depended on the taxa involved and I could not eliminate environmental differences through a short term caging study. Foraging ability of green crab varied in intraspecific competition experiments, with populations from NL and Prince Edward Island dominating longer-established populations from Nova Scotia and New Brunswick. Additional studies excluded claw size as a factor driving these results and behavioral differences likely reflected differences in invasion time and population genetics. Overall, green crab in Placentia Bay appear to be altering community structure of benthic invertebrates through predation and they also appear to indirectly impact native crustaceans through competition.

Environmental data from coastal sediments along natural CO2 gradients at a volcanic vent in Papua New Guinea

Natural CO2 venting systems can mimic conditions that resemble intermediate to high pCO2 levels as predicted for our future oceans. They represent ideal sites to investigate potential long-term effects of ocean acidification on marine life. To test whether microbes are affected by prolonged exposure to pCO2 levels, we examined the composition and diversity of microbial communities in oxic sandy sediments along a natural CO2 gradient. Increasing pCO2 was accompanied by higher bacterial richness and by a strong increase in rare members in both bacterial and archaeal communities. Microbial communities from sites with CO2 concentrations close to today's conditions had different structures than those of sites with elevated CO2 levels. We also observed increasing sequence abundance of several organic matter degrading types of Flavobacteriaceae and Rhodobacteraceae, which paralleled concurrent shifts in benthic cover and enhanced primary productivity. With increasing pCO2, sequences related to bacterial nitrifying organisms such as Nitrosococcus and Nitrospirales decreased, and sequences affiliated to the archaeal ammonia-oxidizing Thaumarchaeota Nitrosopumilus maritimus increased. Our study suggests that microbial community structure and diversity, and likely key ecosystem functions, may be altered in coastal sediments by long-term CO2 exposure to levels predicted for the end of the century.