Nutrient cycling in Alaskan tundra in response to experimental manipulation of growing season length and soil temperature : a climate change scenario

Climate change in the Arctic is predicted to increase plant productivity through decomposition-related enhanced nutrient availability. However, the extent of the increase will depend on whether the increased nutrient availability can be sustained. To address this uncertainty, I assessed the response of plant tissue nutrients, litter decomposition rates, and soil nutrient availability to experimental climate warming manipulations, extended growing season and soil warming, over a 7 year period. Overall, the most consistent effect was the year-to-year variability in measured parameters, probably a result of large differences in weather and time of snowmelt. The results of this study emphasize that although plants of arctic environments are specifically adapted to low nutrient availability, they also posses a suite of traits that help to reduce nutrient losses such as slow growth, low tissue concentrations, and low tissue turnover that result in subtle responses to environmental changes.

Leaf morphology scales multi-annual trends in nutrient cycling and leaf, flower, and fruiting phenology among species in the sub-tropical hardwood forests of the northern Florida Keys

The subtropical hardwood forests of southern Florida are formed by 120 frost-sensitive, broadleaved angiosperm species that range throughout the Caribbean. Previous work on a series of small sized forest component patches of a 20 km2, forest preserve in northern Key Largo indicate that a shift in species composition was associated with a 100 year forest developmental sequence, and this shift was associated with an increasingly evergreen canopy. This document investigates the underlying differences of the biology of trees that live in this habitat, and is specifically focused on the impact of leaf morphology on changing nutrient cycling patterns. Measurements of the area, thickness, dry mass, nutrient content and longevity of several leaves from 3-4 individuals of ten species were conducted in combination with a two-year leaf litter collection and nutrient analysis to determine that species with thicker, denser leaves cycled scarce nutrients up to 2-3 times more efficiently than thin leaved tree species, and the leaf thickness/density index predicts role in forest development in a parallel direction as the index predicts nutrient cycling efficiency. A three year set of observations on the relative abundance of new leaves, flowers and fruits of the same tree species provides an opportunity to evaluate the consequences the leaf morphology/nutrient cycling/forest development relationship to forest habitat quality. Results of the three documents support a mechanistic link between forest development and nutrient cycling, and suggests that older forests are likely to be better habitats based on the availability of valuable forest products like new leaves, flowers, and fruits throughout the year.

Carbon and nitrogen stable isotopic patterns in South Florida coastal ecosystems: Modern and paleoceanographic perspectives

Long term management plans for restoration of natural flow conditions through the Everglades increase the importance of understanding potential nutrient impacts of increased freshwater delivery on coastal biogeochemistry. The present study sought to increase understanding of the coastal marine system of South Florida under modern conditions and through the anthropogenic changes in the last century, on scales ranging from individual nutrient cycle processes to seasonal patterns in organic material (OM) under varying hydrodynamic regime, to century scale analysis of sedimentary records. In all applications, carbon and nitrogen stable isotopic compositions of OM were examined as natural recorders of change and nutrient cycling in the coastal system. ^ High spatial and temporal variability in stable isotopic compositions were observed on all time scales. During a transient phytoplankton bloom, δ 15N values suggested nitrogen fixation as a nutrient source supporting enhanced productivity. Seasonally, particulate organic material (POM) from ten sites along the Florida Reef Tract and in Florida Bay demonstrated variable fluctuations dependent on hydrodynamic setting. Three separate intra-annual patterns were observed, yet statistical differences were observed between groupings of Florida Bay and Atlantic Ocean sites. The POM δ 15N values ranged on a quarterly basis by 7‰, while δ 13C varied by 22‰. From a sediment history perspective, four cores collected from Florida Bay further demonstrated the spatial and temporal variability of the system in isotopic composition of bulk OM over time. Source inputs of OM varied with location, with terrestrial inputs dominating proximal to Everglades freshwater discharge, seagrasses dominating in open estuary cores, and a marine mixture of phytoplankton and seagrass in a core from the boundary zone between Florida Bay and the Gulf of Mexico. Significant shifts in OM geochemistry were observed coincident with anthropogenic events of the 20th century, including railroad and road construction in the Florida Keys and Everglades, and also the extensive drainage changes in Everglades hydrology. The sediment record also preserved evidence of the major hurricanes of the last century, with excursions in geochemical composition coincident with Category 4-5 storms. ^

Epiphyte loads on seagrasses and microphytobenthos abundance are not reliable indicators of nutrient availability in oligotrophic coastal ecosystems

Despite marked gradients in nutrient availability that control the abundance and species composition of seagrasses in south Florida, and the importance of nutrient availability in controlling abundance and composition of epiphytes on seagrasses in other locations, we did not find that epiphyte load on the dominant seagrass, Thalassia testudinum, or that the relative contribution of algal epiphytes to the epiphyte community, was positively correlated with nutrient availability in the water column or the sediment in oligotrophic seagrass beds. Further, the abundance of microphytobenthos, as indicated by Chlorophyll-aconcentration in the sediments, was not directly correlated with concentrations of nutrients in the sediments. Our results suggest that epiphyte and microphytobenthos abundance are not unambiguous indicators of nutrient availability in relatively pristine seagrass environments, and therefore would make poor candidates for indicators of the status and trends of seagrass ecosystems in relatively low-nutrient environments like the Florida Keys.

The Role of the Everglades Mangrove Ecotone Region (EMER) in Regulating Nutrient Cycling and Wetland Productivity in South Florida

The authors summarize the main findings of the Florida Coastal Everglades Long-Term Ecological Research (FCE-LTER) program in the EMER, within the context of the Comprehensive Everglades Restoration Plan (CERP), to understand how regional processes, mediated by water flow, control population and ecosystem dynamics across the EMER landscape. Tree canopies with maximum height <3 m cover 49% of the EMER, particularly in the SE region. These scrub/dwarf mangroves are the result of a combination of low soil phosphorus (P < 59 μg P g dw−1) in the calcareous marl substrate and long hydroperiod. Phosphorus limits the EMER and its freshwater watersheds due to the lack of terrigenous sediment input and the phosphorus-limited nature of the freshwater Everglades. Reduced freshwater delivery over the past 50 years, combined with Everglades compartmentalization and a 10 cm rise in coastal sea level, has led to the landward transgression (1.5 km in 54 years) of the mangrove ecotone. Seasonal variation in freshwater input strongly controls the temporal variation of nitrogen and P exports (99%) from the Everglades to Florida Bay. Rapid changes in nutrient availability and vegetation distribution during the last 50 years show that future ecosystem restoration actions and land use decisions can exert a major influence, similar to sea level rise over the short term, on nutrient cycling and wetland productivity in the EMER.

Carbon and Nitrogen Stable Isotopic Patterns in South Florida Coastal Ecosystems: Modern and Paleoceanographic Perspectives

Long term management plans for restoration of natural flow conditions through the Everglades increase the importance of understanding potential nutrient impacts of increased freshwater delivery on coastal biogeochemistry. The present study sought to increase understanding of the coastal marine system of South Florida under modern conditions and through the anthropogenic changes in the last century, on scales ranging from individual nutrient cycle processes to seasonal patterns in organic material (OM) under varying hydrodynamic regime, to century scale analysis of sedimentary records. In all applications, carbon and nitrogen stable isotopic compositions of OM were examined as natural recorders of change and nutrient cycling in the coastal system. High spatial and temporal variability in stable isotopic compositions were observed on all time scales. During a transient phytoplankton bloom, ä15N values suggested nitrogen fixation as a nutrient source supporting enhanced productivity. Seasonally, particulate organic material (POM) from ten sites along the Florida Reef Tract and in Florida Bay demonstrated variable fluctuations dependent on hydrodynamic setting. Three separate intra-annual patterns were observed, yet statistical differences were observed between groupings of Florida Bay and Atlantic Ocean sites. The POM ä15N values ranged on a quarterly basis by 7‰, while ä13C varied by 22‰. From a sediment history perspective, four cores collected from Florida Bay further demonstrated the spatial and temporal variability of the system in isotopic composition of bulk OM over time. Source inputs of OM varied with location, with terrestrial inputs dominating proximal to Everglades freshwater discharge, seagrasses dominating in open estuary cores, and a marine mixture of phytoplankton and seagrass in a core from the boundary zone between Florida Bay and the Gulf of Mexico. Significant shifts in OM geochemistry were observed coincident with anthropogenic events of the 20th century, including railroad and road construction in the Florida Keys and Everglades, and also the extensive drainage changes in Everglades hydrology. The sediment record also preserved evidence of the major hurricanes of the last century, with excursions in geochemical composition coincident with Category 4-5 storms.

Relative significance of pelagic, benthic and allochthonous organic matter to microbial carbon cycling in a phosphorus-depleted subtropical estuary (Florida Bay)

Heterotrophic bacteria are important decomposers and transformers of primary production and provide an important link between detritus and the aquatic food web. In seagrass ecosystems, much of seagrass primary production is unavailable through direct grazing and must undergo microbial reworking before seagrass production can enter the aquatic food web. The goal of my dissertation research is to understand better the role heterotrophic bacteria play in carbon cycling in seagrass estuaries. My dissertation research focuses on Florida Bay, a seagrass estuary that has experienced recent changes in carbon source availability, which may have altered ecosystem function. My dissertation research investigates the importance of seagrass, algal and/or cyanobacterial, and allochthonous-derived organic matter to heterotrophic bacteria in Florida Bay and helps establish the carbon base of the estuarine food web. ^ A three tiered approach to the study of heterotrophic bacterial carbon cycling and trophic influences in Florida Bay was used: (1) Spatiotemporal observations of environmental parameters (hydrology, nutrients, extracellular enzymes, and microbial abundance, biomass, and production); (2) Microbial grazing experiments under different levels of top-down and bottom-up influence; and (3) Bulk and compound-specific (bacteria-biomarker fatty acid analysis) stable carbon isotope analysis. ^ In Florida Bay, spatiotemporal patterns in microbial extracellular enzyme (also called ectoenzyme) activities indicate that microorganisms hydrolyzed selectively fractions of the estuarine organic matter pool. The microbial community hydrolyzed organic acids, peptides, and phosphate esters and did not use storage and structural carbohydrates. Organic matter use by heterotrophic bacterioplankton in Florida Bay was co-regulated by bottom-up (resource availability) and top-down (grazer mediated) processes. A bacterial carbon budget based on bacterial, epiphytic, and seagrass production indicates that heterotrophic bacterial carbon cycles are supported primarily through epiphytic production with mixing from seagrass production. Stable carbon isotope analysis of bacteria biomarkers and carbon sources in Florida Bay corroborate the results of the bacterial carbon budget. These results support previous studies of aquatic consumers in Florida Bay, indicating that epiphytic/benthic algal and/or cyanobacterial production with mixing from seagrass-derived organic matter is the carbon base of the seagrass estuarine food web. ^

Carbon and nutrient storage in subtropical seagrass meadows: examples from Florida Bay and Shark Bay

Seagrass meadows in Florida Bay and Shark Bay, contain substantial stores of both organic carbon and nutrients. Soils from both systems are predominantly calcium carbonate, with an average of 82.1% CaCO3 in Florida Bay compared to 71.3% in Shark Bay. Soils from Shark Bay had, on average, 21% higher organic carbon content and 35% higher phosphorus content than Florida Bay. Further, soils from Shark Bay had lower mean dry bulk density (0.78 ± 0.01 g mL-1) than those from Florida Bay (0.84 ± 0.02 mg mL-1). The most hypersaline regions of both bays had higher organic carbon content in surficial soils. Profiles of organic carbon and phosphorus from Florida Bay indicate that this system has experienced an increase in P delivery and primary productivity over the last century; in contrast, decreasing organic carbon and phosphorus with depth in the soil profiles in Shark Bay point to a decrease in phosphorus delivery and primary productivity over the last 1000 y. The total ecosystem stocks of stored organic C in Florida Bay averages 163.5 MgCorg ha-1, lower than the average of 243.0 MgCorg ha-1 for Shark Bay; but these values place Shark and Florida Bays among the global hotspots for organic C storage in coastal ecosystems.

Distribution of Diatoms and Development of Diatom-Based Models for Inferring Salinity and Nutrient Concentrations in Florida Bay and Adjacent Coastal Wetlands of South Florida (USA)

The composition and distribution of diatom algae inhabiting estuaries and coasts of the subtropical Americas are poorly documented, especially relative to the central role diatoms play in coastal food webs and to their potential utility as sentinels of environmental change in these threatened ecosystems. Here, we document the distribution of diatoms among the diverse habitat types and long environmental gradients represented by the shallow topographic relief of the South Florida, USA, coastline. A total of 592 species were encountered from 38 freshwater, mangrove, and marine locations in the Everglades wetland and Florida Bay during two seasonal collections, with the highest diversity occurring at sites of high salinity and low water column organic carbon concentration (WTOC). Freshwater, mangrove, and estuarine assemblages were compositionally distinct, but seasonal differences were only detected in mangrove and estuarine sites where solute concentration differed greatly between wet and dry seasons. Epiphytic, planktonic, and sediment assemblages were compositionally similar, implying a high degree of mixing along the shallow, tidal, and storm-prone coast. The relationships between diatom taxa and salinity, water total phosphorus (WTP), water total nitrogen (WTN), and WTOC concentrations were determined and incorporated into weighted averaging partial least squares regression models. Salinity was the most influential variable, resulting in a highly predictive model (r apparent 2  = 0.97, r jackknife 2  = 0.95) that can be used in the future to infer changes in coastal freshwater delivery or sea-level rise in South Florida and compositionally similar environments. Models predicting WTN (r apparent 2  = 0.75, r jackknife 2  = 0.46), WTP (r apparent 2  = 0.75, r jackknife 2  = 0.49), and WTOC (r apparent 2  = 0.79, r jackknife 2  = 0.57) were also strong, suggesting that diatoms can provide reliable inferences of changes in solute delivery to the coastal ecosystem.

Ocean Acidification Outweighs Nutrient Effects in Structuring Seagrass Epiphyte Communities

Developing a framework for assessing interactions between multiple anthropogenic stressors remains an important goal in environmental research. In coastal ecosystems, the relative effects of aspects of global climate change (e.g. CO2 concentrations) and localized stressors (e.g. eutrophication), in combination, have received limited attention. Using a long-term (11 month) field experiment, we examine how epiphyte assemblages in a tropical seagrass meadow respond to factorial manipulations of dissolved carbon dioxide (CO2(aq)) and nutrient enrichment. In situ CO2(aq) manipulations were conducted using clear, open-top chambers, which replicated carbonate parameter forecasts for the year 2100. Nutrient enrichment consisted of monthly additions of slow-release fertilizer, nitrogen (N) and phosphorus (P), to the sediments at rates equivalent to theoretical maximum rates of anthropogenic loading within the region (1.54 g N m−2 d−1 and 0.24 g P m−2 d−1). Epiphyte community structure was assessed on a seasonal basis and revealed declines in the abundance of coralline algae, along with increases in filamentous algae under elevated CO2(aq). Surprisingly, nutrient enrichment had no effect on epiphyte community structure or overall epiphyte loading. Interactions between CO2(aq) and nutrient enrichment were not detected. Furthermore, CO2(aq)-mediated responses in the epiphyte community displayed strong seasonality, suggesting that climate change studies in variable environments should be conducted over extended time-scales. Synthesis. The observed responses indicate that for certain locations, global stressors such as ocean acidification may take precedence over local eutrophication in altering the community structure of seagrass epiphyte assemblages. Given that nutrient-driven algal overgrowth is commonly cited as a widespread cause of seagrass decline, our findings highlight that alternate climate change forces may exert proximate control over epiphyte community structure.

Changing Bacterial Growth Efficiencies across a Natural Nutrient Gradient in an Oligotrophic Estuary

Recent studies have characterized coastal estuarine systems as important components of the global carbon cycle. This study investigated carbon cycling through the microbial loop of Florida Bay by use of bacterial growth efficiency calculations. Bacterial production, bacterial respiration, and other environmental parameters were measured at three sites located along a historic phosphorus-limitation gradient in Florida Bay and compared to a relatively nutrient enriched site in Biscayne Bay. A new method for measuring bacterial respiration in oligotrophic waters involving tracing respiration of 13C-glucose was developed. The results of the study indicate that 13C tracer assays may provide a better means of measuring bacterial respiration in low nutrient environments than traditional dissolved oxygen consumption-based methods due to strong correlations between incubation length and δ13C values. Results also suggest that overall bacterial growth efficiency may be lower at the most nutrient limited sites.

Stable isotopes reveal complex changes in trophic relationships following nutrient addition in a coastal marine ecosystem

Complex links between the top-down and bottomup forces that structure communities can be disrupted by anthropogenic alterations of natural habitats.We used relative abundance and stable isotopes to examine changes in epifaunal food webs in seagrass (Thalassia testudinum) beds following 6 months of experimental nutrient addition at two sites in Florida Bay (USA) with different ambient fertility. At a eutrophic site, nutrient addition did not strongly affect food web structure, but at a nutrient-poor site, enrichment increased the abundances of crustacean epiphyte grazers, and the diets of these grazers became more varied. Benthic grazers did not change in abundance but shifted their diet away from green macroalgae + associated epiphytes and towards an opportunistic seagrass (Halodule wrightii) that occurred only in nutrient addition treatments. Benthic predators did not change in abundance, but their diets were more varied in enriched plots. Food chain length was short and unaffected by site or nutrient treatment, but increased food web complexity in enriched plots was suggested by increasingly mixed diets. Strong bottom-up modifications of food web structure in the nutrient-limited site and the limited top-down influences of grazers on seagrass epiphyte biomass suggest that, in this system, the bottom-up role of nutrient enrichment can have substantial impacts on community structure, trophic relationships, and, ultimately, the productivity values of the ecosystem.

Relating precipitation and water management to nutrient concentrations in the oligotrophic ‘‘upside-down’’ estuaries of the Florida Everglades

We present 8 yr of long-term water quality, climatological, and water management data for 17 locations in Everglades National Park, Florida. Total phosphorus (P) concentration data from freshwater sites (typically ,0.25 mmol L21, or 8 mg L21) indicate the oligotrophic, P-limited nature of this large freshwater–estuarine landscape. Total P concentrations at estuarine sites near the Gulf of Mexico (average ø0.5 m mol L21) demonstrate the marine source for this limiting nutrient. This ‘‘upside down’’ phenomenon, with the limiting nutrient supplied by the ocean and not the land, is a defining characteristic of the Everglade landscape. We present a conceptual model of how the seasonality of precipitation and the management of canal water inputs control the marine P supply, and we hypothesize that seasonal variability in water residence time controls water quality through internal biogeochemical processing. Low freshwater inflows during the dry season increase estuarine residence times, enabling local processes to control nutrient availability and water quality. El Nin˜o–Southern Oscillation (ENSO) events tend to mute the seasonality of rainfall without altering total annual precipitation inputs. The Nin˜o3 ENSO index (which indicates an ENSO event when positive and a La Nin˜a event when negative) was positively correlated with both annual rainfall and the ratio of dry season to wet season precipitation. This ENSO-driven disruption in seasonal rainfall patterns affected salinity patterns and tended to reduce marine inputs of P to Everglades estuaries. ENSO events also decreased dry season residence times, reducing the importance of estuarine nutrient processing. The combination of variable water management activities and interannual differences in precipitation patterns has a strong influence on nutrient and salinity patterns in Everglades estuaries.

Patterns of nutrient exchange in a riverine mangrove forest in the Shark River Estuary, Florida, USA

This study aimed to evaluate tidal and seasonal variations in concentrations and fluxes of nitrogen (NH4 +, NO2+NO3, total nitrogen) and phosphorus (soluble reactive phosphorus, total phosphorus) in a riverine mangrove forest using the flume technique during the dry (May, December 2003) and rainy (October 2003) seasons in the Shark River Estuary, Florida. Tidal water temperatures during the sampling period were on average 29.4 (± 0.4) oC in May and October declining to 20 oC (± 4) in December. Salinity values remained constant in May (28 ± 0.12 PSU), whereas salinity in October and December ranged from 6‒21 PSU and 9‒25 PSU, respectively. Nitrate + nitrite (N+N) and NH4+ concentrations ranged from 0.0 to 3.5 μM and from 0 to 4.8 μM throughout the study period, respectively. Mean TN concentrations in October and December were 39 (±0.8) μM and 37 (±1.5) μM, respectively. SRP and N+N concentrations in the flume increased with higher frequency in flooding tides. TP concentrations ranged between 0.2‒2.9 μM with higher concentrations in the dry season than in the rainy season. Mean concentrations were <1. 5 μM during the sampling period in October (0.75 ± 0.02) and December (0.76 ± 0.01), and were relatively constant in both upstream and downstream locations of the flume. Water residence time in the flume (25 m2) was relatively short for any nutrient exchange to occur between the water column and the forest floor. However, the distinct seasonality in nutrient concentrations in the flume and adjacent tidal creek indicate that the Gulf of Mexico is the main source of SRP and N+N into the mangrove forest.

A preliminary analysis of the correlation of food-web characteristics with hydrology and nutrient gradients in the southern Everglades

We estimated trophic position and carbon source for three consumers (Florida gar, Lepisosteus platyrhincus; eastern mosquitofish, Gambusia holbrooki; and riverine grass shrimp, Palaemonetes paludosus) from 20 sites representing gradients of productivity and hydrological disturbance in the southern Florida Everglades, U.S.A. We characterized gross primary productivity at each site using light/dark bottle incubation and stem density of emergent vascular plants. We also documented nutrient availability as total phosphorus (TP) in floc and periphyton, and the density of small fishes. Hydrological disturbance was characterized as the time since a site was last dried and the average number of days per year the sites were inundated for the previous 10 years. Food-web attributes were estimated in both the wet and dry seasons by analysis of δ15N (trophic position) and δ13C (food-web carbon source) from 702 samples of aquatic consumers. An index of carbon source was derived from a two-member mixing model with Seminole ramshorn snails (Planorbella duryi) as a basal grazing consumer and scuds (amphipods Hyallela azteca) as a basal detritivore. Snails yielded carbon isotopic values similar to green algae and diatoms, while carbon values of scuds were similar to bulk periphyton and floc; carbon isotopic values of cyanobacteria were enriched in C13compared to all consumers examined. A carbon source similar to scuds dominated at all but one study site, and though the relative contribution of scud-like and snail-like carbon sources was variable, there was no evidence that these contributions were a function of abiotic factors or season. Gar consistently displayed the highest estimated trophic position of the consumers studied, with mosquitofish feeding at a slightly lower level, and grass shrimp feeding at the lowest level. Trophic position was not correlated with any nutrient or productivity parameter, but did increase for grass shrimp and mosquitofish as the time following droughts increased. Trophic position of Florida gar was positively correlated with emergent plant stem density.