Patterns of Root Dynamics in Mangrove Forests Along Environmental Gradients in the Florida Coastal Everglades, USA

Patterns of mangrove vegetation in two distinct basins of Florida Coastal Everglades (FCE), Shark River estuary and Taylor River Slough, represent unique opportunities to test hypotheses that root dynamics respond to gradients of resources, regulators, and hydroperiod. We propose that soil total phosphorus (P) gradients in these two coastal basins of FCE cause specific patterns in belowground biomass allocation and net primary productivity that facilitate nutrient acquisition, but also minimize stress from regulators and hydroperiod in flooded soil conditions. Shark River basin has higher P and tidal hydrology with riverine mangroves, in contrast to scrub mangroves of Taylor basin with more permanent flooding and lower P across the coastal landscape. Belowground biomass (0–90 cm) of mangrove sites in Shark River and Taylor River basins ranged from 2317 to 4673 g m-2, with the highest contribution (62–85%) of roots in the shallow root zone (0–45 cm) compared to the deeper root zone (45–90 cm). Total root productivity did not vary significantly among sites and ranged from 407 to 643 g m-2 y-1. Root production in the shallow root zone accounted for 57–78% of total production. Root turnover rates ranged from 0.04 to 0.60 y-1 and consistently decreased as the root size class distribution increased from fine to coarse roots, indicating differences in root longevity. Fine root biomass was negatively correlated with soil P density and frequency of inundation, whereas fine root turnover decreased with increasing soil N:P ratios. Lower P availability in Taylor River basin relative to Shark River basin, along with higher regulator and hydroperiod stress, confirms our hypothesis that interactions of stress from resource limitation and long duration of hydroperiod account for higher fine root biomass along with lower fine root production and turnover. Because fine root production and organic matter accumulation are the primary processes controlling soil formation and accretion in scrub mangrove forests, root dynamics in the P-limited carbonate ecosystem of south Florida have a major controlling role as to how mangroves respond to future impacts of sealevel rise.

Water source utilization and foliar nutrient status differs between upland and flooded plant communities in wetland tree islands

Tree islands in the Everglades wetlands are centers of biodiversity and targets of restoration, yet little is known about the pattern of water source utilization by the constituent woody plant communities: upland hammocks and flooded swamp forests. Two potential water sources exist: (1) entrapped rainwater in the vadose zone of the organic soil (referred to as upland soil water), that becomes enriched in phosphorus, and (2) phosphorus-poor groundwater/surface water (referred to as regional water). Using natural stable isotope abundance as a tracer, we observed that hammock plants used upland soil water in the wet season and shifted to regional water uptake in the dry season, while swamp forest plants used regional water throughout the year. Consistent with the previously observed phosphorus concentrations of the two water sources, hammock plants had a greater annual mean foliar phosphorus concentration over swamp forest plants, thereby supporting the idea that tree island hammocks are islands of high phosphorus concentrations in the oligotrophic Everglades. Foliar nitrogen levels in swamp forest plants were higher than those of hammock plants. Linking water sources with foliar nutrient concentrations can indicate nutrient sources and periods of nutrient uptake, thereby linking hydrology with the nutrient regimes of different plant communities in wetland ecosystems. Our results are consistent with the hypotheses that (1) over long periods, upland tree island communities incrementally increase their nutrient concentration by incorporating marsh nutrients through transpiration seasonally, and (2) small differences in micro-topography in a wetland ecosystem can lead to large differences in water and nutrient cycles.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Abstract Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

Vegetation dynamics and their implications for the management of wetlands in the lowlands of Nepal

The maintenance of species richness is often a priority in the management of nature reserves, where consumptive use of resources is generally prohibited. The purpose of this research was to improve management by understanding the vegetation dynamics in the lowlands of Nepal. The objectives were to determine vegetation associations in relation to environments and human-induced disturbances that affect vegetation dynamics on floodplains, where upstream barrages had altered flooding patterns, and consumptive use of plant resources was influencing natural processes. Floodplain vegetation in relation to physical environments and disturbances were studied along transects, perpendicular to the course of the Mahakali River in the western Terai, Nepal. Forest structural changes were studied for three years in ten plots. A randomized split-block experiment with nine burning and grazing treatments was performed in seasonally flooded grasslands. A semi-structured questionnaire was used to assess people's socio-economic status, natural resource use patterns and conservation attitudes. ^ Elevation, soil organic matter, nitrogen, percentage of sand and grazing intensity were significant in delineating herbaceous vegetation assemblages, whereas elevation and livestock grazing were significant in defining forest type boundaries. On the floodplain islands, highly grazed Dalbergia sissoo-Acacia catechu forests were devoid of understory woody vegetation, but the lightly grazed D. sissoo-mixed forests had a well-developed second canopy layer, comprising woody species other than D. sissoo and A. catechu. In grasslands, species richness and biomass production were highest at intermediate disturbance level represented by the lightly grazed and ungrazed early-burned treatments. Ethnicity, education and resource use patterns were important in influencing conservation attitudes. A succession towards the mixed forests would occur in D. sissoo-dominated floodplain forests, where dams and barrages reduce flooding and associated fluvial processes, and if livestock grazing is stopped, as occasionally suggested by nature conservationists. In seasonally flooded grasslands, early burning with moderate grazing would enhance the species diversity and productivity. There is a need to implement a participatory integrated wetland management plan, to include community development, education and off farm income generation, to assure participatory conservation and management of wetlands in Nepal. ^

Soil carbon dynamics in primary and secondary tropical forests in Colombia

Soils play a central role in the dynamics of biospheric carbon and in climate change. They contain the largest carbon stock of terrestrial ecosystems and return to the atmosphere a significant proportion of carbon fixed by photosynthesis. Soils of tropical forests are tremendously important in the carbon cycle because they receive the largest organic matter inputs, they have the largest respiration rates, and they are among the largest carbon reservoirs among world soils. This research assesses the main components of the soil carbon dynamics in primary (PF) and secondary (SF) tropical forests in Colombia. I evaluated the production, stocks, and decomposition rates of aboveground detritus as well as the stocks, growth, mortality, and decomposition of fine roots in these two forest types. Soil carbon outputs were evaluated as total soil, heterotrophic, and root respiration. The stocks of soil organic carbon down to 4 m deep in these two cover types and in degraded pastures (PAS) were also evaluated. ^ Soil inputs of organic carbon from above and belowground sources were lower in SF than in PF. Litterfall in SF was 58% and production of fine root detritus was 60% of that in PF. When production of woody detritus and palm fronds was considered, the difference between these forest types was even larger. However, outputs of mineral carbon through heterotrophic soil respiration were similar; in SF they equaled 97% of those in PF. As a result, soil carbon balance was positive in PF and negative in SF. Despite that soil carbon balances suggest that soils of SF are losing carbon, soil carbon stocks of SF were higher than of degraded pastures, suggesting that they have already started to recover soil carbon stocks lost under degraded pastures. This discrepancy can be partially explained by the effect of drier conditions on heterotrophic soil respiration as a consequence of a moderate El Niño event during the period of soil respiration measurements. The positive carbon balance in soils of PF despite the El Niño event, suggests that soils of PF accumulated about 664 Kg C ha−1 yr−1. Therefore, soil carbon dynamics mainly depended on successional status of vegetation and on climatic conditions. ^

The successional dynamics of lightning-initiated canopy gaps in the mangrove forests of Shark River, Everglades National Park, USA

Gap succession is a significant determinant of structure and development in most forest communities. Lightning strikes are an important source of canopy gaps in the mangrove forest of Everglades National Park. I investigated the successional dynamics of lightning-initiated canopy gaps to determine their influence on forest stand structure of the mixed mangrove forests ( Rhizophora mangle, Laguncularia racemosa, and Avicennia germinans ) of the Shark River. I measured gap size, gap shape, light environment, soil characteristics, woody debris, and fiddler crab abundance. I additionally measured the vegetative composition in a chronosequences of gap successional stages (new, recruiting, and growing gaps). I recorded survivorship, recruitment, growth and soil elevation dynamics within a subset of new and growing gaps. I determined the relationship between intact forest soil elevation and site hydrology in order to interpret the effects of lightning disturbance on soil elevation dynamics. ^ Gap size averaged 289 ± 20 m2 (± 1SE) and light transmittance decreased exponentially as gaps filled with saplings. Fine woody debris was highest in recruiting gaps. Soil strength was lower in the gaps than in the forest. The abundance of large and medium fiddler crab burrows increased linearly with total seedling abundance. Soil surface elevation declined in newly formed lightning gaps; this loss was due to a combination of superficial erosion (8.5 mm) and subsidence (60.9 mm). A distinct two-cohort recruitment pattern was evident in the seedling/sapling surveys, suggesting a partitioning of the succession between individuals present before and after lightning strike. In new gaps, the seedling recruitment rate was twice as high as in forest and the sapling population increased. At the growing gap stage, R. mangle seedling mortality was 10 times greater and sapling mortality was 13 times greater than recruitment. Growing gaps had reduced seedling stem elongation, sapling growth and adult growth. However, a few individuals (R. mangle saplings) were able to recruit into the adult life stage. In conclusion, the high density of R. mangle seedlings and saplings imply that lightning strike disturbances in these mangrove forests favor their recruitment over that of A. germinans and L. racemosa. ^

Ecosystem responses to hydrologic change and mechanisms of nitrogen sequestration in seasonally flooded tree islands of the southern Everglades

In the Florida Everglades, tree islands are conspicuous heterogeneous elements in a complex wetland landscape. I investigated the effects of increased freshwater flow in southern Everglades seasonally flooded tree islands, and characterized biogeochemical interactions among tree islands and the marsh landscape matrix, specifically examining hydrologic flows of nitrogen (N), and landscape N sequestration capacity. I utilized ecological trajectories of key ecosystem variables to differentiate effects of increased sheetflow and hydroperiod. I utilized stable isotope analyses and nutrient content of tree island ecosystem components to test the hypothesis that key processes in tree island nitrogen cycling would favor ecosystem N sequestration. I combined estimates of tree island ecosystem N standing stocks and fluxes, soil and litter N transformation rates, and hydrologic inputs of N to quantify the net sequestration of N by a seasonally flooded tree island. ^ Results show that increased freshwater flow to seasonally flooded tree islands promoted ecosystem oligotrophy, whereas reduced flows allowed some plant species to cycle P less efficiently. As oligotrophy is a defining parameter of Everglades wetlands, and likely promotes belowground production and peat development, reintroducing freshwater flow from an upstream canal had a favorable effect on ecosystem dynamics of tree islands in the study area. Important factors influencing the stable isotopic composition of nitrogen and carbon were: (1) a contribution to soil N by soil invertebrates, animal excrement, and microbes, (2) a possible NO3 source from an upstream canal and an "open" ecosystem N cycle, and (3) greater availability of phosphorus in tree islands relative to the marsh landscape, suggesting that tree island N cycling favors N sequestration. Hydrologic sources of N were dominated by surface water loads of NO3- and NH 4+, and an important soil N transformation promoting the net loss of surface water DIN was nitrate immobilization associated with soils and surficial leaf litter. The net inorganic N sequestration capacity of a seasonally flooded tree island was 50 g yr-1 m -2. Thus, tree islands likely have an important function in landscape sequestration of inorganic N, and may reduce significant anthropogenic N loads to downstream coastal systems. ^

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.

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.

Chilling damage in a changing climate in coastal landscapes of the subtropical zone: a case study from south Florida

Extensive portions of the southern Everglades are characterized by series of elongated, raised peat ridges and tree islands oriented parallel to the predominant flow direction, separated by intervening sloughs. Tall herbs or woody species are associated with higher elevations and shorter emergent or floating species are associated with lower elevations. The organic soils in this “Ridge-and-Slough” landscape have been stable over millennia in many locations, but degrade over decades under altered hydrologic conditions. We examined soil, pore water, and leaf phosphorus (P) and nitrogen (N) distributions in six Ridge and Slough communities in Shark Slough, Everglades National Park. We found P enrichment to increase and N to decrease monotonically along a gradient from the most persistently flooded sloughs to rarely flooded ridge environments, with the most dramatic change associated with the transition from marsh to forest. Leaf N:P ratios indicated that the marsh communities were strongly P-limited, while data from several forest types suggested either N-limitation or co-limitation by N and P. Ground water stage in forests exhibited a daytime decrease and partial nighttime recovery during periods of surface exposure. The recovery phase suggested re-supply from adjacent flooded marshes or the underlying aquifer, and a strong hydrologic connection between ridge and slough. We therefore developed a simple steady-state model to explore a mechanism by which a phosphorus conveyor belt driven by both evapotranspiration and the regional flow gradient can contribute to the characteristic Ridge and Slough pattern. The model demonstrated that evapotranspiration sinks at higher elevations can draw in low concentration marsh waters, raising local soil and water P concentrations. Focusing of flow and nutrients at the evapotranspiration zone is not strong enough to overcome the regional gradient entirely, allowing the nutrient to spread downstream and creating an elongated concentration plume in the direction of flow. Our analyses suggest that autogenic processes involving the effects of initially small differences in topography, via their interactions with hydrology and nutrient availability, can produce persistent physiographic patterns in the organic sediments of the Everglades.

Woody Debris in the Mangrove Forests of South Florida

Woody debris is abundant in hurricane-impacted forests. With a major hurricane affecting South Florida mangroves approximately every 20 yr, carbon storage and nutrient retention may be influenced greatly by woody debris dynamics. In addition, woody debris can influence seedling regeneration in mangrove swamps by trapping propagules and enhancing seedling growth potential. Here, we report on line-intercept woody debris surveys conducted in mangrove wetlands of South Florida 9–10 yr after the passage of Hurricane Andrew. The total volume of woody debris for all sites combined was estimated at 67 m3/ha and varied from 13 to 181 m3/ha depending upon differences in forest height, proximity to the storm, and maximum estimated wind velocities. Large volumes of woody debris were found in the eyewall region of the hurricane, with a volume of 132 m3/ha and a projected woody debris biomass of approximately 36 t/ha. Approximately half of the woody debris biomass averaged across all sites was associated as small twigs and branches (fine woody debris), since coarse woody debris >7.5 cm felled during Hurricane Andrew was fairly well decomposed. Much of the small debris is likely to be associated with post-hurricane forest dynamics. Hurricanes are responsible for large amounts of damage to mangrove ecosystems, and components of associated downed wood may provide a relative index of disturbance for mangrove forests. Here, we suggest that a fine:coarse woody debris ratio ≤0.5 is suggestive of a recent disturbance in mangrove wetlands, although additional research is needed to corroborate such findings.

Sea level rise and South Florida coastal forests

Coastal ecosystems lie at the forefront of sea level rise. We posit that before the onset of actual inundation, sea level rise will influence the species composition of coastal hardwood hammocks and buttonwood (Conocarpus erectus L.) forests of the Everglades National Park based on tolerance to drought and salinity. Precipitation is the major water source in coastal hammocks and is stored in the soil vadose zone, but vadose water will diminish with the rising water table as a consequence of sea level rise, thereby subjecting plants to salt water stress. A model is used to demonstrate that the constraining effect of salinity on transpiration limits the distribution of freshwater-dependent communities. Field data collected in hardwood hammocks and coastal buttonwood forests over 11 years show that halophytes have replaced glycophytes. We establish that sea level rise threatens 21 rare coastal species in Everglades National Park and estimate the relative risk to each species using basic life history and population traits. We review salinity conditions in the estuarine region over 1999–2009 and associate wide variability in the extent of the annual seawater intrusion to variation in freshwater inflows and precipitation. We also examine species composition in coastal and inland hammocks in connection with distance from the coast, depth to water table, and groundwater salinity. Though this study focuses on coastal forests and rare species of South Florida, it has implications for coastal forests threatened by saltwater intrusion across the globe.

Mapping Height and Biomass of Mangrove Forests in Everglades National Park with SRTM Elevation Data

We produced a landscape scale map of mean tree height in mangrove forests in Everglades National Park (ENP) using the elevation data from the Shuttle Radar Topography Mission (SRTM). The SRTM data was calibrated using airborne lidar data and a high resolution USGS digital elevation model (DEM). The resulting mangrove height map has a mean tree height error of 2.0 m (RMSE) over a pixel of 30 m. In addition, we used field data to derive a relationship between mean forest stand height and biomass in order to map the spatial distribution of standing biomass of mangroves for the entire National Park. The estimation showed that most of the mangrove standing biomass in the ENP resides in intermediate- height mangrove stands around 8 m. We estimated the total mangrove standing biomass in ENP to be 5.6 X 109 kg.

Sediment accretion and organic carbon burial relative to sea-level rise and storm events in two mangrove forests in Everglades National Park

The goal of this investigation was to examine how sediment accretion and organic carbon (OC) burial rates in mangrove forests respond to climate change. Specifically, will the accretion rates keep pace with sea-level rise, and what is the source and fate of OC in the system? Mass accumulation, accretion and OC burial rates were determined via 210Pb dating (i.e. 100 year time scale) on sediment cores collected from two mangrove forest sites within Everglades National Park, Florida (USA). Enhanced mass accumulation, accretion and OC burial rates were found in an upper layer that corresponded to a well-documented storm surge deposit. Accretion rates were 5.9 and 6.5 mm yr− 1 within the storm deposit compared to overall rates of 2.5 and 3.6 mm yr− 1. These rates were found to be matching or exceeding average sea-level rise reported for Key West, Florida. Organic carbon burial rates were 260 and 393 g m− 2 yr− 1 within the storm deposit compared to 151 and 168 g m− 2 yr− 1 overall burial rates. The overall rates are similar to global estimates for OC burial in marine wetlands. With tropical storms being a frequent occurrence in this region the resulting storm surge deposits are an important mechanism for maintaining both overall accretion and OC burial rates. Enhanced OC burial rates within the storm deposit could be due to an increase in productivity created from higher concentrations of phosphorus within storm-delivered sediments and/or from the deposition of allochthonous OC. Climate change-amplified storms and sea-level rise could damage mangrove forests, exposing previously buried OC to oxidation and contribute to increasing atmospheric CO2 concentrations. However, the processes described here provide a mechanism whereby oxidation of OC would be limited and the overall OC reservoir maintained within the mangrove forest sediments.