Controls on Ecosystem Carbon Dioxide Exchange in Short- and Long-Hydroperiod Florida Everglades Freshwater Marshes

Although freshwater wetlands are among the most productive ecosystems on Earth, little is known of carbon dioxide (CO2) exchange in low latitude wetlands. The Everglades is an extensive, oligotrophic wetland in south Florida characterized by short- and long-hydroperiod marshes. Chamber-based CO2 exchange measurements were made to compare the marshes and examine the roles of primary producers, seasonality, and environmental drivers in determining exchange rates. Low rates of CO2 exchange were observed in both marshes with net ecosystem production reaching maxima of 3.77 and 4.28 μmol CO2 m−2 s−1 in short- and long-hydroperiod marshes, respectively. Fluxes of CO2 were affected by seasonality only in the short-hydroperiod marsh, where flux rates were significantly lower in the wet season than in the dry season. Emergent macrophytes dominated fluxes at both sites, though this was not the case for the short-hydroperiod marsh in the wet season. Water depth, a factor partly under human control, significantly affected gross ecosystem production at the short-hydroperiod marsh. As Everglades ecosystem restoration proceeds, leading to deeper water and longer hydroperiods, productivity in short-hydroperiod marshes will likely be more negatively affected than in long-hydroperiod marshes. The Everglades stand in contrast to many freshwater wetlands because of ecosystem-wide low productivity rates.

Phosphorus enrichment effects on interactions among the ecosystem components in a long-hydroperiod oligotrophic marsh in Everglades National Park

Hydrology and a history of oligotrophy unite the massive landscape comprising freshwater marsh in Everglades National Park. With restoration of water flow to the Everglades, phosphorus (P) enrichment, both from agricultural and domestic sources, may increase nutrient load to the marsh ecosystem. Previous research of P enrichment of Everglades soil, periphyton, and macrophytes revealed each of these ecosystem components responds to increased P loads with increased production and nutrient content. Interactions among these ecosystem components and how P affects the magnitude and direction of interaction are poorly understood and are the focus of my research. Here I present results of a two-year, two-factor experiment of P enrichment and manipulation in Everglades National Park. I quantified biomass, nutrient content, and production for periphyton and macrophyes and found macrophyte removal drives change in nutrient content, biomass, and production of periphyton. Periphyton removal did not appear to control macrophyte dynamics. Soil chemical and physical characteristics were explained primarily by site differences but there was an enrichment effect of soil porewater nitrite + nitrate, nitrite, and soluble reactive phosphorus. Flocculent materials production and depth were significantly affected by macrophyte removal where depth and production were significantly greater with the no macrophyte treatment. The dominant macrophyte of the marsh, Eleocharis cellulosa, increased more in the unenriched marsh than in the enriched marsh. The combination of these findings suggests that dynamics in floc and periphyton are controlled primarily by the presence of periphyton and that this relationship is significantly affected by low-level P enrichment. These results may be valuable in their application to both managers and policy makers who are involved in the Everglades restoration process. ^

Short and long hydroperiod Everglades periphyton mats: Community characterization and experimental hydroperiod manipulation

Hydroperiod, or the distribution, duration and timing of flooding affects both plant and animal distributions. The Florida Everglades is currently undergoing restoration that will result in altered hydroperiods. This study was conducted in Everglades National Park to document the variability in periphyton community structure and function between long and short hydroperiod Everglades marshes. Periphyton is an important primary producer and important food resource in the Everglades. Periphyton is also involved in marl soil formation and nutrient cycling. Although periphyton is an important component of the Everglades landscape, little is known about periphyton structural-functional variation between hydroperiods. ^ For this study diatoms, as well as fresh algae slides of diatoms, cyanobacteria and green algae were identified and enumerated. Short verse long hydroperiod soil and water column nutrients were compared. Short and long hydroperiod algal periphyton mat productivity rates were compared using BOD incubations. Experimental manipulations were performed to determine the effects of desiccation duration and rewetting on periphyton productivity, community structure, and nutrient flux. ^ Variation in periphyton community structure was significantly greater between hydroperiods than within hydroperiods. Short and long hydroperiod periphyton mats have the same algal species, it is the distribution and abundance that varies between hydroperiods. Long hydroperiod mats have greater diatom abundance while short hydroperiod mats have greater relative filamentous cyanobacterial abundance. ^ Long hydroperiod mats had greater net primary production (npp) than short hydroperiod mats. Short hydroperiod mats respond to rewetting more rapidly than do long hydroperiod mats. Dry short hydroperiod mats became net primary producers within 24 hours of rehydration. Increasing desiccation duration led to greater cyanobacterial abundance in long hydroperiod mats and decreased diatom abundance in both long and short hydroperiod mats. ^

Eddy covariance and meteorological tower in a long-hydroperiod Everglades marsh during the height of the wet season, 2008 (at SRS-2), Shark River Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Long-hydroperiod Everglades marsh, wet season, 2008 (near SRS-2), Shark River Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Eddy covariance tower in long-hydroperiod Everglades marsh in mid-dry season, 2009 (at SRS-2), Shark River Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Long-hydroperiod Everglades marsh under extremely dry conditions, late dry season, 2009 (near SRS-2), Shark River Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Seasonal Fish Dispersal in Ephemeral Wetlands of the Florida Everglades

We hypothesized that fishes in short-hydroperiod wetlands display pulses in activity tied to seasonal flooding and drying, with relatively low activity during intervening periods. To evaluate this hypothesis, sampling devices that funnel fish into traps (drift fences) were used to investigate fish movement across the Everglades, U.S.A. Samples were collected at six sites in the Rocky Glades, a seasonally flooded karstic habitat located on the southeastern edge of the Everglades. Four species that display distinct recovery patterns following drought in long-hydroperiod wetlands were studied: eastern mosquitofish (Gambusia holbrooki) and flagfish (Jordanella floridae) (rapid recovery); and bluefin killifish (Lucania goodei) and least killifish (Heterandria formosa) (slow recovery). Consistent with our hypothesized conceptual model, fishes increased movement soon after flooding (immigration period) and just before drying (emigration period), but decreased activity in the intervening foraging period. We also found that eastern mosquitofish and flagfish arrived earlier and showed stronger responses to hydrological variation than either least killifish or bluefin killifish. We concluded that these fishes actively colonize and escape ephemeral wetlands in response to flooding and drying, and display species-specific differences related to flooding and drying that reflect differences in dispersal ability. These results have important implications for Everglades fish metacommunity dynamics.

Vegetation:environment relationships and water management in Shark Slough, Everglades National Park

The hydrologic regime of Shark Slough, the most extensive long hydroperiod marsh in Everglades National Park, is largely controlled by the location, volume, and timing of water delivered to it through several control structures from Water Conservation Areas north of the Park. Where natural or anthropogenic barriers to water flow are present, water management practices in this highly regulated system may result in an uneven distribution of water in the marsh, which may impact regional vegetation patterns. In this paper, we use data from 569 sampling locations along five cross-Slough transects to examine regional vegetation distribution, and to test and describe the association of marsh vegetation with several hydrologic and edaphic parameters. Analysis of vegetation:environment relationships yielded estimates of both mean and variance in soil depth, as well as annual hydroperiod, mean water depth, and 30-day maximum water depth within each cover type during the 1990’s. We found that rank abundances of the three major marsh cover types (Tall Sawgrass, Sparse Sawgrass, and Spikerush Marsh) were identical in all portions of Shark Slough, but regional trends in the relative abundance of individual communities were present. Analysis also indicated clear and consistent differences in the hydrologic regime of three marsh cover types, with hydroperiod and water depths increasing in the order Tall Sawgrass , Sparse Sawgrass , Spikerush Marsh. In contrast, soil depth decreased in the same order. Locally, these differences were quite subtle; within a management unit of Shark Slough, mean annual values for the two water depth parameters varied less than 15 cm among types, and hydroperiods varied by 65 days or less. More significantly, regional variation in hydrology equaled or exceeded the variation attributable to cover type within a small area. For instance, estimated hydroperiods for Tall Sawgrass in Northern Shark Slough were longer than for Spikerush Marsh in any of the other regions. Although some of this regional variation may reflect a natural gradient within the Slough, a large proportion is the result of compartmentalization due to current water management practices within the marsh.We conclude that hydroperiod or water depth are the most important influences on vegetation within management units, and attribute larger scale differences in vegetation pattern to the interactions among soil development, hydrology and fire regime in this pivotal portion of Everglades.

Crayfish assemblage shifts in a large drought-prone wetland: the roles of hydrology and competition

1. Faster growing, larger and/or more aggressive crayfish species are predicted to dominate permanent waterbodies. We tested this prediction using a 9 year dataset for two species of crayfish (Procambarus alleni and Procambarus fallax) co-existing in a sub-tropical flowing slough in southern Florida. Using a series of laboratory and mesocosm experiments we also compared life history traits and performance of the respective species to test mechanisms that could explain dominance shifts in the local crayfish assemblages. 2. Over the 9-year period, P. alleni densities were the greatest in shallower, shorterhydroperiod areas bordering the slough, while P. fallax densities were higher in deeper, longer-hydroperiod central areas. These areas were separated by 0.8–2 km of continuous wetland with no apparent barriers to movement between them. 3. Density of P. fallax was not strongly affected by any measures of hydrological variation, while P. alleni density increased with more severe drought conditions. Following the strongest droughts, P. alleni colonized areas in the centre of the slough where they had been absent or scarce in wetter years. 4. We conducted experiments to compare growth rates, drought tolerance, and competitive dominance of these species. P. alleni survived drought conditions better, had higher growth rates, and was the dominant competitor for space and food. While drought probably limits P. fallax in the drier slough habitats, neither drought sensitivity nor interspecific competition with P. fallax can explain decreases of P. alleni with wetter conditions. 5. Our results indicate that a competition-colonization tradeoff cannot explain the crayfish compositional dynamics in this wetland because P. alleni is both the best competitor and the best at surviving in and colonizing areas with the strongest droughts. Future attention should focus on the potential for selective effects of predators that co-vary with hydrology. 6. The traits (large size, fast growth, competitive dominance) exhibited by P. alleni, which is absent in long-hydroperiod wetlands, are those exhibited by dominant crayfish in permanent lakes and streams containing fish. Although these traits make crayfish less vulnerable to fish in some lakes and streams, life-history models of community structure across permanence gradients suggest the opposite traits should be favoured for co-existence with fish.

Comparative study of periphyton community structure in long and short-hydroperiod Everglades marshes

The Florida Everglades is a mosaic of short and long-hydroperiod marshes that differ in the depth, duration, and timing of inundation. Algae are important primary producers in widespread Everglades’ periphyton mats, but relationships of algal production and community structure to hydrologic variability are poorly understood. We quantified differences in algal biomass and community structure between periphyton mats in 5 short and 6 long-hydroperiod marshes in Everglades National Park (ENP) in October 2000. We related differences to water depth and total phosphorus (TP) concentration in the water, periphyton and soils. Long and short-hydroperiod marshes differed in water depth (73 cm vs. 13 cm), periphyton TP concentrations (172μg g−1 vs. 107 μg g−1, respectively) and soil TP (284 μg g−1 vs. 145 μg g−1). Periphyton was abundant in both marshes, with short-hydroperiod sites having greater biomass than long-hydroperiod sites (2936 vs. 575 grams ash-free dry mass m−2). A total of 156 algal taxa were identified and separated into diatom (68 species from 21 genera) and “soft algae” (88 non-diatom species from 47 genera) categories for further analyses. Although diatom total abundance was greater in long-hydroperiod mats, diatom species richness was significantly greater in short- hydroperiod periphyton mats (62 vs. 47 diatom taxa). Soft algal species richness was greater in long-hydroperiod sites (81 vs. 67 soft algae taxa). Relative abundances of individual taxa were significantly different among the two site types, with soft algal distributions being driven by water depth, and diatom distributions by water depth and TP concentration in the water and periphyton. Periphyton communities differ between short and long-hydroperiod marshes, but because they share many taxa, alterations in hydroperiod could rapidly

Interactions between nutrient availability and hydroperiod shape macroinvertebrate communities in Florida Everglades marshes

Hydroperiod and nutrient status are known to influence aquatic communities in wetlands, but their joint effects are not well explored. I sampled floating periphyton mat and flocculent detritus (floc) infaunal communities using 6-cm diameter cores at short- and long-hydroperiod and constantly inundated sites across a range of phosphorus (P) availability (total phosphorus in soil, floc and periphyton). Differences in community structure between periphyton and floc microhabitats were greater than any variation attributable to hydroperiod, P availability, or other spatial factors. Multivariate analyses indicated community structure of benthic-floc infauna was driven by hydroperiod, although crowding (no. g−1 AFDM) of individual taxa showed no consistent responses to hydroperiod or P availability. In contrast, community structure of periphyton mat infauna was driven by P availability, while densities of mat infauna (no. m−2) were most influenced by hydroperiod (+correlations). Crowding of mat infauna increased significantly with P availability in short-hydroperiod marshes, but was constant across the P gradient in long-hydroperiod marshes. Increased abundance of floating-periphyton mat infauna with P availability at short-hydroperiod sites may result from a release from predation by small fish. Community structure and density were not different between long-hydroperiod and constantly inundated sites. These results have implications for the use of macroinvertebrates as indicators of water quality in wetlands and suggest the substrate sampled can influence interpretation of ecological responses observed in these communities.

Seasonal differences in the CO2 exchange of a short-hydroperiod Florida Everglades marsh

Although wetlands are among the world's most productive ecosystems, little is known of long-term CO2 exchange in tropical and subtropical wetlands. The Everglades is a highly managed wetlands complex occupying >6000 km2 in south Florida. This ecosystem is oligotrophic, but extremely high rates of productivity have been previously reported. To evaluate CO2 exchange and its response to seasonality (dry vs. wet season) in the Everglades, an eddy covariance tower was established in a short-hydroperiod marl marsh. Rates of net ecosystem exchange and ecosystem respiration were small year-round and declined in the wet season relative to the dry season. Inundation reduced macrophyte CO2 uptake, substantially limiting gross ecosystem production. While light and air temperature exerted the primary controls on net ecosystem exchange and ecosystem respiration in the dry season, inundation weakened these relationships. The ecosystem shifted from a CO2 sink in the dry season to a CO2 source in the wet season; however, the marsh was a small carbon sink on an annual basis. Net ecosystem production, ecosystem respiration, and gross ecosystem production were −49.9, 446.1 and 496.0 g C m−2 year−1, respectively. Unexpectedly low CO2 flux rates and annual production distinguish the Everglades from many other wetlands. Nonetheless, impending changes in water management are likely to alter the CO2 balance of this wetland and may increase the source strength of these extensive short-hydroperiod wetlands.

Fish Community Responses to the Combined Effects of Decreased Hydroperiod and Nonnative Fish Invasions in a Karst Wetland: Are Everglades Solution Holes Sinks for Native Fishes?

Short-hydroperiod Everglades wetlands have been disproportionately affected by reductions in freshwater inflows, land conversion and biotic invasions. Severe hydroperiod reductions in these habitats, including the Rocky Glades, coupled with proximity to canals that act as sources of invasions, may limit their ability to support high levels of aquatic production. We examined whether karst solution holes function as dry-down refuges for fishes, providing a source of marsh colonists upon reflooding, by tracking fish abundance, nonnative composition, and survival in solution holes throughout the dry season. We paired field surveys with an in situ nonnative predation experiment that tested the effects of predation by the recent invader, African jewelfish (Hemichromis letourneuxi) on native fishes. Over the 3 years surveyed, a large number of the solution holes dried before the onset of the wet season, while those retaining water had low survivorship and were dominated by nonnatives. In the experiment, mortality of eastern mosquitofish (Gambusia holbrooki) in the presence of African jewelfish was greater than that associated with deteriorating water quality. Under current water management, findings suggest that solution holes are largely sinks for native fishes, given the high frequency of drydown, extensive period of fish residence, and predation by nonnative fishes.

A influência dos fragmentos florestais na dinâmica de populações de anuros no noroeste do Estado de São Paulo

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)