Macroinvertebrate community response to eutrophication in an oligotrophic wetland: An in situ mesocosm experiment

Eutrophication from anthropogenic nutrient enrichment is a primary threat to the oligotrophic freshwater marshes of southern Florida. Macrophyte and periphyton response to increased phosphorus (P) has been well documented in both correlative and experimental studies, but the response of consumer communities remains poorly understood, especially in southern marl prairies. We conducted a P-loading experiment in in situ mesocosms in Taylor Slough, Everglades National Park, and examined the response of macroinvertebrate communities. Mesocosms at two sites were loaded weekly with P at four levels: control (0 g P/m2/yr), low (0.2 g P/m2/yr), intermediate (0.8 g P/m2/yr), and high (3.2 g P/m2/ yr). After ∼2 yrs of P-loading, macroinvertebrates were sampled using periphyton mat and benthic floc cores. Densities of macroinvertebrate taxa (no./g AFDM) were two to 16 times higher in periphyton mats than benthic floc. Periphyton biomass decreased with enrichment at one site, and periphyton was absent from many intermediate and all high P treatments at both sites. Total macroinvertebrate density in periphyton mats increased with intermediate P loads, driven primarily by chironomids and nematodes. Conversely, total macroinvertebrate density in benthic floc decreased with enrichment, driven primarily by loss of chironomids and ceratopogonids (Dasyhelea). This study suggests that macroinvertebrate density increases with enrichment until periphyton mats are lost, after which it decreases, and mat infauna fail to move into benthic substrates in response to mat loss. These results were noted at nutrient levels too low to yield anoxia, and we believe that the decrease of macroinvertebrate density resulted from a loss of habitat. This work illustrates the importance of periphyton mats as habitat for macroinvertebrates in the Everglades. This study also indicates that in this system, macroinvertebrate sampling should be designed to target periphyton mats or conducted with special attention to inclusion of substrates relative to their coverage.

Molecular characterization of dissolved organic matter in freshwater wetlands of the Florida Everglades

In this study, the molecular composition of dissolved organic matter (DOM), collected from wetlands of the Southern Everglades, was examined using a variety of analytical techniques in order to characterize its sources and transformation in the environment. The methods applied for the characterization of DOM included fluorescence spectroscopy, solid state 13C CPMAS NMR spectroscopy, and pyrolysis-GC/MS. The relative abundance of protein-like components and carbohydrates increased from the canal site to more remote freshwater marsh sites suggesting that significant amounts of non-humic DOM are autochthonously produced within the freshwater marshes, and are not exclusively introduced through canal inputs. Such in situ DOM production is important when considering how DOM from canals is processed and transported to downstream estuaries of Florida Bay.

Subtroprical wetland fish assemblages and changing salinity regimes: Implications for everglades restoration

During the 1960s, water management practices resulted in the conversion of the wetlands that fringe northeastern Florida Bay (USA) from freshwater/oligohaline herbaceous marshes to dwarf red mangrove forests. Coincident with this conversion were several ecological changes to Florida Bay’s fauna, including reductions in the abundances of top trophic-level consumers: piscivorous fishes, alligators, crocodiles, and wading birds. Because these taxa rely on a common forage base of small demersal fishes, food stress has been implicated as playing a role in their respective declines. In the present study, we monitored the demersal fishes seasonally at six sites over an 8-year time period. During monitoring, extremely high rainfall conditions occurred over a 3.5-year period leading to salinity regimes that can be viewed as “windows” to the area’s natural past and future restored states. In this paper, we: (1) examine the changes in fish communities over the 8-year study period and relate them to measured changes in salinity; (2) make comparisons among marine, brackish and freshwater demersal fish communities in terms of species composition, density, and biomass; and (3) discuss several implications of our findings in light of the intended and unintended water management changes that are planned or underway as part of Everglades restoration. Results suggest the reduction in freshwater flow to Florida Bay over the last several decades has reduced demersal fish populations, and thus prey availability for apex consumers in the coastal wetlands compared to the pre-drainage inferred standard. Furthermore, greater discharge of freshwater toward Florida Bay may result in the re-establishment of pre-1960s fauna, including a more robust demersal-fish community that should prompt increases in populations of several important predatory species.

Phosphorus cycling and partitioning in an oligotrophic Everglades wetland ecosystem: a radioisotope tracing study

1. Our goal was to quantify short-term phosphorus (P) partitioning and identify the ecosystem components important to P cycling in wetland ecosystems. To do this, we added P radiotracer to oligotrophic, P-limited Everglades marshes. 32PO4 was added to the water column in six 1-m2 enclosed mesocosms located in long-hydroperiod marshes of Shark River Slough, Everglades National Park. Ecosystem components were then repeatedly sampled over 18 days. 2. Water column particulates (>0.45 μm) incorporated radiotracer within the first minute after dosing and stored 95–99% of total water column 32P activity throughout the study. Soluble (<0.45 μm) 32P in the water column, in contrast, was always <5% of the 32P in surface water. Periphyton, both floating and attached to emergent macrophytes, had the highest specific activity of 32P (Bq g−131P) among the different ecosystem components. Fish and aquatic macroinvertebrates also had high affinity for P, whereas emergent macrophytes, soil and flocculent detrital organic matter (floc) had the lowest specific activities of radiotracer. 3. Within the calcareous, floating periphyton mats, 81% of the initial 32P uptake was associated with Ca, but most of this 32P entered and remained within the organic pool (Ca-associated = 14% of total) after 1 day. In the floc layer, 32P rapidly entered the microbial pool and the labile fraction was negligible for most of the study. 4. Budgeting of the radiotracer indicated that 32P moved from particulates in the water column to periphyton and floc and then to the floc and soil over the course of the 18 day incubations. Floc (35% of total) and soil (27%) dominated 32P storage after 18 days, with floating periphyton (12%) and surface water (10%) holding smaller proportions of total ecosystem 32P. 5. To summarise, oligotrophic Everglades marshes exhibited rapid uptake and retention of labile 32P. Components dominated by microbes appear to control short-term P cycling in this oligotrophic ecosystem.

Phosphorus budgets in Everglades wetland ecosystems: the effects of hydrology and nutrient enrichment

The Florida Everglades is a naturally oligotrophic hydroscape that has experienced large changes in ecosystem structure and function as the result of increased anthropogenic phosphorus (P) loading and hydrologic changes. We present whole-ecosystem models of P cycling for Everglades wetlands with differing hydrology and P enrichment with the goal of synthesizing existing information into ecosystem P budgets. Budgets were developed for deeper water oligotrophic wet prairie/slough (‘Slough’), shallower water oligotrophic Cladium jamaicense (‘Cladium’), partially enriched C. jamaicense/Typha spp. mixture (‘Cladium/Typha’), and enriched Typha spp. (‘Typha’) marshes. The majority of ecosystem P was stored in the soil in all four ecosystem types, with the flocculent detrital organic matter (floc) layer at the bottom of the water column storing the next largest proportion of ecosystem P pools. However, most P cycling involved ecosystem components in the water column (periphyton, floc, and consumers) in deeper water, oligotrophic Slough marsh. Fluxes of P associated with macrophytes were more important in the shallower water, oligotrophic Cladium marsh. The two oligotrophic ecosystem types had similar total ecosystem P stocks and cycling rates, and low rates of P cycling associated with soils. Phosphorus flux rates cannot be estimated for ecosystem components residing in the water column in Cladium/Typha or Typha marshes due to insufficient data. Enrichment caused a large increase in the importance of macrophytes to P cycling in Everglades wetlands. The flux of P from soil to the water column, via roots to live aboveground tissues to macrophyte detritus, increased from 0.03 and 0.2 g P m−2 yr−1 in oligotrophic Slough and Cladium marsh, respectively, to 1.1 g P m−2 yr−1 in partially enriched Cladium/Typha, and 1.6 g P m−2 yr−1 in enriched Typha marsh. This macrophyte translocation P flux represents a large source of internal eutrophication to surface waters in P-enriched areas of the Everglades.

Disturbance regime and limits on benefits of refuge use for fishes in a fluctuating hydroscape

Refuge habitats increase survival rate and recovery time of populations experiencing environmental disturbance, but limits on the ability of refuges to buffer communities are poorly understood. We hypothesized that importance of refuges in preventing population declines and alteration in community structure has a non-linear relationship with severity of disturbance. In the Florida Everglades, alligator ponds are used as refuge habitat by fishes during seasonal drying of marsh habitats. Using an 11-year record of hydrological conditions and fish abundance in 10 marshes and 34 alligator ponds from two regions of the Everglades, we sought to characterize patterns of refuge use and temporal dynamics of fish abundance and community structure across changing intensity, duration, and frequency of drought disturbance. Abundance in alligator ponds was positively related to refuge size, distance from alternative refugia (e.g. canals), and abundance in surrounding marsh prior to hydrologic disturbance. Variables negatively related to abundance in alligator ponds included water level in surrounding marsh and abundance of disturbance-tolerant species. Refuge community structure did not differ between regions because the same subset of species in both regions used alligator ponds during droughts. When time between disturbances was short, fish abundance declined in marshes, and in the region with the most spatially extensive pattern of disturbance, community structure was altered in both marshes and alligator ponds because of an increased proportion of species more resistant to disturbance. These changes in community structure were associated with increases in both duration and frequency of hydrologic disturbance. Use of refuge habitat had a modal relationship with severity of disturbance regime. Spatial patterns of response suggest that decline in refuge use was because of decreased effectiveness of refuge habitat in reducing mortality and providing sufficient time for recovery for fish communities experiencing reduced time between disturbance events.

Does variation in movement tactics and trophic interactions among American alligators create habitat linkages?

1. Highly mobile top predators are hypothesized to spatially and/or temporally link disparate habitats through the combination of their movement and feeding patterns, but recent studies suggest that individual specialization in habitat use and feeding could keep habitats compartmentalized. 2.  We used passive acoustic telemetry and stable isotope analysis to investigate whether specialization in movement and feeding patterns of American alligators (Alligator mississippiensis) in an oligotrophic subtropical estuary created habitat linkages between marine and estuarine/freshwater food webs. 3.  Individual alligators adopted one of the three relatively distinct movement tactics that were linked to variation in diets. Fifty-six per cent of alligators regularly travelled from the upstream (freshwater/mid-estuary) areas into the downstream (marine-influenced) areas where salinities exceed those typically tolerated by alligators. Thirty-one per cent of the alligators made regular trips from the mid-estuarine habitat into the upstream habitat; 13% remained in the mid-estuary zone year-round. 4.  Stable isotopic analysis indicated that, unlike individuals remaining in the mid-estuary and upstream zones, alligators that used the downstream zone fed at least partially from marine food webs and likely moved to access higher prey abundance at the expense of salt stress. Therefore, ‘commuting’ alligators may link marine food webs with those of the estuary and marshes in the coastal Everglades and create an upstream vector for allochthonous nutrient inputs into the estuary. 5.  This study lends further support to the hypothesis that large-bodied highly mobile predators faced with trade-offs are likely to exhibit individual specialization leading to habitat linkages, rather than compartmentalization. However, the conditions under which this scenario occurs require further investigation.

Comparison of Belowground Biomass in C3- and C4-Dominated Mixed Communities in a Chesapeake Bay Brackish Marsh

Belowground biomass is a critical factor regulating ecosystem functions of coastal marshes, including soil organic matter (SOM) accumulation and the ability of these systems to keep pace with sea-level rise. Nevertheless, belowground biomass responses to environmental and vegetation changes have been given little emphasis marsh studies. Here we present a method using stable carbon isotopes and color to identify root and rhizomes of Schoenoplectus americanus (Pers.) Volk. ex Schinz and R. Keller (C3) and Spartina patens (Ait.) Muhl. (C4) occurring in C3− and C4-dominated communities in a Chesapeake Bay brackish marsh. The functional significance of the biomass classes we identified is underscored by differences in their chemistry, depth profiles, and variation in biomass and profiles relative to abiotic and biotic factors. C3 rhizomes had the lowest concentrations of cellulose (29.19%) and lignin (14.43%) and the lowest C:N (46.97) and lignin:N (0.16) ratios. We distinguished two types of C3 roots, and of these, the dark red C3 roots had anomalously high C:N (195.35) and lignin:N (1.14) ratios, compared with other root and rhizome classes examined here and with previously published values. The C4-dominated community had significantly greater belowground biomass (4119.1 g m−2) than the C3-dominated community (3256.9 g m−2), due to greater total root biomass and a 3.6-fold higher C3-root:rhizome ratio in the C4-dominated community. C3 rhizomes were distributed significantly shallower in the C4-dominated community, while C3 roots were significantly deeper. Variability in C3 rhizome depth distributions was explained primarily by C4 biomass, and C3 roots were explained primarily by water table height. Our results suggest that belowground biomass in this system is sensitive to slight variations in water table height (across an 8 cm range), and that the reduced overlap between C3 and C4 root profiles in the C4-dominated community may account for the greater total root biomass observed in that community. Given that future elevated atmospheric CO2 and accelerated sea-level rise are likely to increase C3 abundance in Atlantic and Gulf coast marshes, investigations that quantify how patterns of C3 and C4 belowground biomass respond to environmental and biological factors stand to improve our understanding of ecosystem-wide impacts of global changes on coastal wetlands.

Ecosystem structure, nutrient dynamics, and hydrologic relationships in tree islands of the southern Everglades, Florida, USA

Tree islands are an important structural component of many graminoid-dominated wetlands because they increase ecological complexity in the landscape. Tree island area has been drastically reduced with hydrologic modifications within the Everglades ecosystem, yet still little is known about the ecosystem ecology of Everglades tree islands. As part of an ongoing study to investigate the effects of hydrologic restoration on short hydroperiod marshes of the southern Everglades, we report an ecosystem characterization of seasonally flooded tree islands relative to locations described by variation in freshwater flow (i.e. locally enhanced freshwater flow by levee removal). We quantified: (1) forest structure, litterfall production, nutrient utilization, soil dynamics, and hydrologic properties of six tree islands and (2) soil and surface water physico-chemical properties of adjacent marshes. Tree islands efficiently utilized both phosphorus and nitrogen, but indices of nutrient-use efficiency indicated stronger P than N limitation. Tree islands were distinct in structure and biogeochemical properties from the surrounding marsh, maintaining higher organically bound P and N, but lower inorganic N. Annual variation resulting in increased hydroperiod and lower wet season water levels not only increased nitrogen use by tree species and decreased N:P values of the dominant plant species (Chrysobalanus icaco), but also increased soil pH and decreased soil temperature. When compared with other forested wetlands, these Everglades tree islands were among the most nutrient efficient, likely a function of nutrient immobilization in soils and the calcium carbonate bedrock. Tree islands of our study area are defined by: (1) unique biogeochemical properties when compared with adjacent short hydroperiod marshes and other forested wetlands and (2) an intricate relationship with marsh hydrology. As such, they may play an important and disproportionate role in nutrient and carbon cycling in Everglades wetlands. With the loss of tree islands that has occurred with the degradation of the Everglades system, these landscape processes may have been altered. With this baseline dataset, we have established a long-term ecosystem-scale experiment to follow the ecosystem trajectory of seasonally flooded tree islands in response to hydrologic restoration of the southern Everglades.

Maintaining tree islands in the Florida Everglades: nutrient redistribution is the key

The Florida Everglades is an oligotrophic wetland system with tree islands as one of its most prominent landscape features. Total soil phosphorus concentrations on tree islands can be 6 to 100 times greater than phosphorus levels in the surrounding marshes and sloughs, making tree islands nutrient hotspots. Several mechanisms are believed to redistribute phosphorus to tree islands: subsurface water flows generated by evapotranspiration of trees, higher deposition rates of dry fallout, deposition of guano by birds and other animals, groundwater upwelling, and bedrock mineralization by tree exudates. A conceptual model is proposed, in which the focused redistribution of limiting nutrients, especially phosphorus, onto tree islands controls their maintenance and expansion. Because of increased primary production and peat accretion rates, the redistribution of phosphorus can result in an increase in both tree island elevation and size. Human changes to hydrology have greatly decreased the number and size of tree islands in parts of the Everglades. The proposed model suggests that the preservation of existing tree islands, and ultimately of the Everglades landscape, requires the maintenance of these phosphorus redistribution mechanisms.

Estimating Global “Blue Carbon” Emissions from Conversion and Degradation of Vegetated Coastal Ecosystems

Recent attention has focused on the high rates of annual carbon sequestration in vegetated coastal ecosystems—marshes, mangroves, and seagrasses—that may be lost with habitat destruction (‘conversion’). Relatively unappreciated, however, is that conversion of these coastal ecosystems also impacts very large pools of previously-sequestered carbon. Residing mostly in sediments, this ‘blue carbon’ can be released to the atmosphere when these ecosystems are converted or degraded. Here we provide the first global estimates of this impact and evaluate its economic implications. Combining the best available data on global area, land-use conversion rates, and near-surface carbon stocks in each of the three ecosystems, using an uncertainty-propagation approach, we estimate that 0.15–1.02 Pg (billion tons) of carbon dioxide are being released annually, several times higher than previous estimates that account only for lost sequestration. These emissions are equivalent to 3–19% of those from deforestation globally, and result in economic damages of $US 6–42 billion annually. The largest sources of uncertainty in these estimates stems from limited certitude in global area and rates of land-use conversion, but research is also needed on the fates of ecosystem carbon upon conversion. Currently, carbon emissions from the conversion of vegetated coastal ecosystems are not included in emissions accounting or carbon market protocols, but this analysis suggests they may be disproportionally important to both. Although the relevant science supporting these initial estimates will need to be refined in coming years, it is clear that policies encouraging the sustainable management of coastal ecosystems could significantly reduce carbon emissions from the land-use sector, in addition to sustaining the well-recognized ecosystem services of coastal habitats.

Relating Freshwater Flow with Estuarine Water Quality in the Southern Everglades Mangrove Ecotone

Florida Bay is more saline than it was historically, and reduced freshwater flows may lead to more phosphorus inputs to the mangrove ecotone from the marine end-member. This is important given plans to restore freshwater flow into eastern Florida Bay. We investigated the relationships between salinity, nutrients, and hydrologic variables in the mangrove ecotone of Taylor Slough. We expected that total phosphorus (TP) would increase with salinity, reflecting a downstream marine source, while total nitrogen (TN) would increase with flow in the mangrove ecotone. Despite expectations of increased flows improving the ecological health of lower Taylor Slough and Florida Bay, total nitrogen (TN) and total phosphorus (TP) dynamics may shift in response to new conditions of flow and salinity as well as organic carbon, N, and P availability. Our results showed that TP concentrations are more discharge-driven while TN is more variable and potentially derived from different sources along the flow path from the freshwater Everglades marshes to Florida Bay. Increased flow of freshwater through Taylor Slough will likely decrease TP concentrations in this historically oligotrophic and P-limited ecosystem. However, more studies along the mangrove ecotone is needed to understand how increased flows will affect nitrogen dynamics relative to phosphorus.

No Free Lunch: Displaced Marsh Consumers Regulate a Prey Subsidy to an Estuarine Consumer

Pulse subsidies account for a substantial proportion of resource availability in many systems, having persistent and cascading effects on consumer population dynamics, and energy flow within and across ecosystem boundaries. Although the importance of pulsed resource subsidies is well-established, the mechanisms that regulate resource fluxes across ecosystem boundaries are not well understood. The aim of our study was to determine the extent that marsh consumers regulated a marsh prey subsidy to estuarine consumers in the oligohaline reaches of an Everglades estuary. We characterized a marsh pulsed subsidy of cyprinodontoid, invertebrate and sunfish prey that move into the upper estuary from adjacent drying marshes. In response to the prey pulse, we examined the numerical, fitness and dietary responses of three focal consumers in the upper estuary; two marsh species (largemouth bass and bowfin) that accompanied the subsidy as a result of marsh drying, and one estuarine consumer (snook). At the onset of marsh drying and the prey subsidy, estuarine consumers switched diets to consume the larger marsh prey (sunfishes), while bass and bowfin maintained similar diets (cyprinodontoids and invertebrates respectively) than pre and post subsidy. From the consumption of this subsidy, bass (marsh species) and snook (estuarine species) exhibited fitness gains while bowfin did not. Although both marsh and estuarine consumers benefitted from the subsidy, we found evidence that freshwater consumers shunted some of the subsidy away from snook. Of the prey sampled in consumer stomachs, 41% of marsh prey biomass was eaten by marsh consumers, while 59% was consumed by the estuarine consumer. We conclude that the amount of the marsh prey available to estuarine consumers may be greater in the absence of marsh consumers, thus the magnitude of the prey subsidy could depend on the dynamics of the marsh consumers from donor communities.

Effect of Water Management on Interannual Variation in Bulk Soil Properties from the Eastern Coastal Everglades

We examined interannual variation in soil properties from wetlands occurring in adjacent drainage basins from the southeastern Everglades. Triplicate 10-cm soil cores were collected, homogenized, and analyzed during the wet season 2006–2010 from five freshwater sawgrass wetland marshes and three estuarine mangrove forests. Soil bulk density from the Taylor Slough basin ranged from 0.15 gm-cm−3 to 0.5 gm-cm−3, was higher than from the Panhandle basin every year, and generally increased throughout the study period. Organic matter as a percent loss on ignition ranged from 7 % to 12 % from freshwater marshes and from 13 % to 56 % from estuarine mangroves. Extractable iron in soils was similar among drainage basins and wetland types, typically ranging from 0.6 to 2.0 g Fe kg−1. In contrast, inorganic sulfur was on average over four times higher from estuarine soils relative to freshwater, and was positively correlated with soil organic matter. Finally total soil phosphorus (P) was lower in freshwater soils relative to estuarine soils (84 ± 5 versus 326 ± 32 mg P kg−1). Total P from the freshwater marshes in the Panhandle basin rose throughout the study period from 54.7 ± 8.4 to 107 ± 17 mg P kg−1, a possible outcome of differences in water management between drainage basins.

Spatial and Temporal Variability of Dissolved Organic Matter Quantity and Composition in an Oilgotrophic Subtropical Coastal Wetland

Dissolved organic matter (DOM) is an essential component of the carbon cycle and a critical driver in controlling variety of biogeochemical and ecological processes in wetlands. The quality of this DOM as it relates to composition and reactivity is directly related to its sources and may vary on temporal and spatial scales. However, large scale, long-term studies of DOM dynamics in wetlands are still scarce in the literature. Here we present a multi-year DOM characterization study for monthly surface water samples collected at 14 sampling stations along two transects within the greater Everglades, a subtropical, oligotrophic, coastal freshwater wetland-mangrove-estuarine ecosystem. In an attempt to assess quantitative and qualitative variations of DOM on both spatial and temporal scales, we determined dissolved organic carbon (DOC) values and DOM optical properties, respectively. DOM quality was assessed using, excitation emission matrix (EEM) fluorescence coupled with parallel factor analysis (PARAFAC). Variations of the PARAFAC components abundance and composition were clearly observed on spatial and seasonal scales. Dry versus wet season DOC concentrations were affected by dry-down and re-wetting processes in the freshwater marshes, while DOM compositional features were controlled by soil and higher plant versus periphyton sources respectively. Peat-soil based freshwater marsh sites could be clearly differentiated from marl-soil based sites based on EEM–PARAFAC data. Freshwater marsh DOM was enriched in higher plant and soil-derived humic-like compounds, compared to estuarine sites which were more controlled by algae- and microbial-derived inputs. DOM from fringe mangrove sites could be differentiated between tidally influenced sites and sites exposed to long inundation periods. As such coastal estuarine sites were significantly controlled by hydrology, while DOM dynamics in Florida Bay were seasonally driven by both primary productivity and hydrology. This study exemplifies the application of long term optical properties monitoring as an effective technique to investigate DOM dynamics in aquatic ecosystems. The work presented here also serves as a pre-restoration condition dataset for DOM in the context of the Comprehensive Everglades Restoration Plan (CERP).