A Site Description of the CARICOMP Mangrove, Seagrass and Coral Reef Sites in Bocas del Toro, Panama

Bocas del Toro is located in the western region of the Republic of Panama. It is part of a province of approximately 8917 km(2) with an estimated 68% of its area covered by tropical rainforest. The area receives 2870 mm/year of rainfall. The dry and rainy seasons are not clearly defined. There are two periods each of low and high rainfall, March and September-October, and July and December, respectively. Mangrove forests, seagrass meadows and coral reefs are vast, covering large areas in the shallow waters surrounding the islands of the archipelago and along the mainland coast. The CARICOMP sites were established in 1998-99 and are periodically monitored following Level I protocol. Herein we describe the sites in a regional context and present the baseline data for each site. This paper fulfills the requirements of the formal site description for CARICOMP monitoring sites.

A review of the potential role of tumour-promoting compounds produced by Lyngbya majuscula in marine turtle fibropapillomatosis

Harmful algal blooms (HABs) have increased in abundance and severity in recent decades. Whereas the implications for human impacts and intoxication resulting from blooms have been extensively studied, the ecological implications of these microalgae are less well understood. Many HAB species produce biologically active, secondary metabolites and the fate of these toxins through the foodweb is generally not well understood unless it culminates in extensive fish mortalities or human poisonings. This review focusses on one HAB species, the cyanobacterium Lyngbya majuscula, and presents a hypothetical role for its involvement in fibro-papillornatosis (FP), a neoplastic disease of marine turtles. FP is expressed as benign tumours that grow both internally and externally on marine turtles, preventing vision, movement and organ function. The aetiology of FP is currently not conclusively understood, but virus material has been associated with tumours and previous studies have suggested a role for naturally produced tumour promoters. In this review, we present a hypothesis regarding the involvement of L. majuscula in FP, either through direct intoxication and action of tumour-promoting compounds or indirectly by causing seagrass loss and compromised immune function, thus leaving the turtles more susceptible to disease.

Effects of Lyngbya majuscula (Cyanophycea) blooms on sediment nutrients and meiofaunal assemblages in seagrass beds in Moreton Bay, Australia

Blooms of Lyngbya majuscula have been increasingly recorded in the waters of Moreton Bay, on the south-east coast of Queensland, Australia. The influences of these blooms on sediment infauna and the implications for sediment biogeochemical processes was studied. Sediment samples were taken from Moreton Bay banks during and after the bloom season. The deposition of L. majuscula seems to be responsible for the higher total Kjedahl nitrogen (TKN) concentrations measured during the bloom period. Total organic carbon (TOC) concentrations did not change. Lyngbya majuscula blooms had a marked influence on the meiobenthos. Nematodes, copepods and polychaetes were the most abundant groups of meiofauna, and the bloom produced a decrease in the abundance and a change in the sediment depth distribution of these organisms. The distribution of nematodes, copepods and polychaetes in sediment became shallower. Further, the bloom did not affect the abundance and distribution of polychaetes as strongly as it did copepods and nematodes. The changes observed in the distribution of meiofauna in the sediment during the bloom period indicate that L. majuscula produces oxygen depletion in sediments, and that different fauna seem to be affected to different degrees.

Effects of physical disturbance on infaunal and epifaunal assemblages in subtropical, intertidal seagrass beds

We assessed the impact of large-scale commercial and recreational harvesting of polychaete worms Marphysa spp. on macrobenthic assemblages in a subtropical estuary in Queensland, Australia, by examining: (1) the spatial extent of harvesting activities and the rate of recovery of the seagrass habitat over an 18 to 20 mo period; (2) the recovery of infauna in and around commercial pits of known age; (3) the indirect effects of physical disturbance from trampling and deposition of sediments during harvesting on epibenthos in areas adjacent to commercial and recreational pits; (4) impacts of potential indirect effects through manipulative experimentation. Harvesting caused a loss of seagrass, changes to the topography and compaction of the sediments associated with the creation of walls around commercial pits, and the deposition of rubble dug from within the pit. The walls and rubble were still evident after 1.8 to 20 mo, but comprised only a small proportion of the total area on the intertidal banks. There was a shift from an intertidal area dominated by Zostera capricorni to one with a mixture of Z. capricorni, Halophila spp. and Halodule uninervis, but there was no overall decline in the biomass of seagrass in these areas. There were distinct impacts from harvesting on the abundance of benthic infauna, especially amphipods, polychaetes and gastropods, and these effects were still detectable after 4 mo of potential recovery. After 12 me, there were no detectable differences in the abundances of these infauna between dug areas and reference areas, which suggested that infauna had recovered from impacts of harvesting; however, an extensive bloom of toxic fireweed Lyngbya majsucula may have masked any remaining impacts. There were no detectable impacts of harvesting on epifauna living in the seagrass immediately around commercial or recreational pits.

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.

Biogeochemical effects of iron availability on primary producers in a shallow marine carbonate environment

We completed a synoptic survey of iron, phosphorus, and sulfur concentrations in shallow marine carbonate sediments from south Florida. Total extracted iron concentrations typically were 50 μmol g-1 dry weight (DW) and tended to decrease away from the Florida mainland, whereas total extracted phosphorus concentrations mostly were 10 μmol g-1 DW and tended to decrease from west to east across Florida Bay. Concentrations of reduced sulfur compounds, up to 40 μmol g-1 DW, tended to covary with sediment iron concentrations, suggesting that sulfide mineral formation was iron-limited. An index of iron availability derived from sediment data was negatively correlated with chlorophyll a concentrations in surface waters, demonstrating the close coupling of sediment-water column processes. Eight months after applying a surface layer of iron oxide granules to experimental plots, sediment iron, phosphorus, and sulfur were elevated to a depth of 10 cm relative to control plots. Biomass of the seagrass Thalassia testudinum was not different between control and iron addition plots, but individual shoot growth rates were significantly higher in experimental plots after 8 months. Although the iron content of leaf tissues was significantly higher from iron addition plots, no difference in phosphorus content of T. testudinum leaves was observed. Iron addition altered plant exposure to free sulfide, documented by a significantly higher δ34S of leaf tissue from experimental plots relative to controls. Iron as a buffer to toxic sulfides may promote individual shoot growth, but phosphorus availability to plants still appears to limit production in carbonate sediments.

Tropical seagrass-associated macroalgae distributions and trends relative to water quality

Tropical coastal marine ecosystems including mangroves, seagrass beds and coral reef communities are undergoing intense degradation in response to natural and human disturbances, therefore, understanding the causes and mechanisms present challenges for scientist and managers. In order to protect our marine resources, determining the effects of nutrient loads on these coastal systems has become a key management goal. Data from monitoring programs were used to detect trends of macroalgae abundances and develop correlations with nutrient availability, as well as forecast potential responses of the communities monitored. Using eight years of data (1996–2003) from complementary but independent monitoring programs in seagrass beds and water quality of the Florida Keys National Marine Sanctuary (FKNMS), we: (1) described the distribution and abundance of macroalgae groups; (2) analyzed the status and spatiotemporal trends of macroalgae groups; and (3) explored the connection between water quality and the macroalgae distribution in the FKNMS. In the seagrass beds of the FKNMS calcareous green algae were the dominant macroalgae group followed by the red group; brown and calcareous red algae were present but in lower abundance. Spatiotemporal patterns of the macroalgae groups were analyzed with a non-linear regression model of the abundance data. For the period of record, all macroalgae groups increased in abundance (Abi) at most sites, with calcareous green algae increasing the most. Calcareous green algae and red algae exhibited seasonal pattern with peak abundances (Φi) mainly in summer for calcareous green and mainly in winter for red. Macroalgae Abi and long-term trend (mi) were correlated in a distinctive way with water quality parameters. Both the Abi and mi of calcareous green algae had positive correlations with NO3−, NO2−, total nitrogen (TN) and total organic carbon (TOC). Red algae Abi had a positive correlation with NO2−, TN, total phosphorus and TOC, and the mi in red algae was positively correlated with N:P. In contrast brown and calcareous red algae Abi had negative correlations with N:P. These results suggest that calcareous green algae and red algae are responding mainly to increases in N availability, a process that is happening in inshore sites. A combination of spatially variable factors such as local current patterns, nutrient sources, and habitat characteristics result in a complex array of the macroalgae community in the seagrass beds of the FKNMS.

The Impact of Hurricane Georges on Soft-Bottom, Back Reef Communitites: Site- and Species-Specific Effects in South Florida Seagrass Beds

Seagrass beds are the dominant benthic marine communities in the back reef environment of the Florida Keys. At a network of 30 permanent monitoring stations in this back reef environment, the seagrass Thalassia testudinum Banks & Soland. ex Koenig was the most common marine macrophyte, but the seagrasses Syringodium fi liforme Kuetz., and Halodule wrightii Aschers., as well as many taxa of macroalgae, were also commonly encountered. The calcareous green macroalgae, especially Halimeda spp. and Penicillus spp., were the most common macroalgae. The passage of Hurricane Georges on September 25, 1998 caused an immediate loss of 3% of the density of T. testudinum, compared to 19% of the S. fi liforme and 24% of the calcareous green algae. The seagrass beds at three of the stations were completely obliterated by the storm. Stations that had little to moderate sediment deposition recovered from the storm within 1 yr, while the station buried by 50 cm of sediment and the two stations that experienced substantial erosion had recovered very little during the 3 yrs after the storm. Early colonizers to these severely disturbed sites were calcareous green algae. Hurricanes may increase benthic macrophyte diversity by creating disturbed patches with the landscape, but moderate storm disturbance may actually reduce macrophyte diversity by removing the early successional species from mixed-species seagrass beds.

Seagrass ecosystems as a globally significant carbon stock

The protection of organic carbon stored in forests is considered as an important method for mitigating climate change. Like terrestrial ecosystems, coastal ecosystems store large amounts of carbon, and there are initiatives to protect these ‘blue carbon’ stores. Organic carbon stocks in tidal salt marshes and mangroves have been estimated, but uncertainties in the stores of seagrass meadows—some of the most productive ecosystems on Earth—hinder the application of marine carbon conservation schemes. Here, we compile published and unpublished measurements of the organic carbon content of living seagrass biomass and underlying soils in 946 distinct seagrass meadows across the globe. Using only data from sites for which full inventories exist, we estimate that, globally, seagrass ecosystems could store as much as 19.9 Pg organic carbon; according to a more conservative approach, in which we incorporate more data from surface soils and depth-dependent declines in soil carbon stocks, we estimate that the seagrass carbon pool lies between 4.2 and 8.4 Pg carbon. We estimate that present rates of seagrass loss could result in the release of up to 299 Tg carbon per year, assuming that all of the organic carbon in seagrass biomass and the top metre of soils is remineralized.

Responses of seagrass communities to fertilization along a gradient of relative availability of nitrogen and phosphorus in a carbonate environment

Patterns of relative nutrient availability in south Florida suggest spatial differences regarding the importance of nitrogen (N) and phosphorus (P) to benthic primary producers. We did a 14-month in situ fertilization experiment to test predictions of N and P limitation in the subtropical nearshore marine waters of the upper Florida Keys. Six sites were divided into two groups (nearshore, offshore) representing the endpoints of an N: P stoichiometric gradient. Twenty-four plots were established at each site with six replicates of each treatment (1N, 1P, 1N1P, control), for a total of 144 experimental plots. The responses of benthic communities to N and P enrichment varied appreciably between nearshore and offshore habitats. Offshore seagrass beds were strongly limited by nitrogen, and nearshore beds were affected by nitrogen and phosphorus. Nutrient addition at offshore sites increased the length and aboveground standing crop of the two seagrasses, Thalassia testudinum and Syringodium filiforme, and growth rates of T. testudinum. Nutrient addition at nearshore sites increased the relative abundance of macroalgae, epiphytes, and sediment microalgae. N limitation of seagrass in this carbonate system was clearly demonstrated. However, added phosphorus was retained in the system more effectively than N, suggesting that phosphorus might have important long-term effects on these benthic communities. The observed species-specific responses to nutrient enrichment underscores the need to monitor all primary producers when addressing questions of nutrient limitation and eutrophication in seagrass communities.

The effects of carbon dioxide fertilization on the ecology of tropical seagrass communities

Increasing atmospheric CO2 concentrations associated with climate change will likely influence a wide variety of ecosystems. Terrestrial research has examined the effects of increasing CO2 concentrations on the functionality of plant systems; with studies ranging in scale from the short-term responses of individual leaves, to long-term ecological responses of complete forests. While terrestrial plants have received much attention, studies on the responses of marine plants (seagrasses) to increased CO 2(aq) concentrations remain relatively sparse, with most research limited to small-scale, ex situ experimentation. Furthermore, few studies have attempted to address similarities between terrestrial and seagrass responses to increases in CO2(aq). The goals of this dissertation are to expand the scope of marine climate change research, and examine how the tropical seagrass, Thalassia testudinum responds to increasing CO 2(aq)concentrations over multiple spatial and temporal scales. ^ Manipulative laboratory and field experimentation reveal that, similar to terrestrial plants, seagrasses strongly respond to increases in CO 2(aq) concentrations. Using a novel field technique, in situ field manipulations show that over short time scales, seagrasses respond to elevated CO2(aq) by increasing leaf photosynthetic rates and the production of soluble carbohydrates. Declines in leaf nutrient (nitrogen and phosphorus) content were additionally detected, paralleling responses from terrestrial systems. Over long time scales, seagrasses increase total above- and belowground biomass with elevated CO2(aq), suggesting that, similar to terrestrial research, pervasive increases in atmospheric and oceanic CO2(aq) concentrations stand to influence the productivity and functionality of these systems. Furthermore, field experiments reveal that seagrass epiphytes, which comprise an important component of seagrass ecosystems, additionally respond to increased CO2(aq) with strong declines in calcified taxa and increases in fleshy taxa. ^ Together, this work demonstrates that increasing CO2(aq) concentrations will alter the functionality of seagrass ecosystems by increasing plant productivity and shifting the composition of the epiphyte community. These results have implications for future rates of carbon storage and sediment production within these widely distributed systems.^

Herbivore and nutrient impact on primary producer assemblages in a tropical marine environment

Globally, human populations are increasing and coastal ecosystems are becoming increasingly impacted by anthropogenic stressors. As eutrophication and exploitation of coastal resources increases, primary producer response to these drivers becomes a key indicator of ecosystem stability. Despite the importance of monitoring primary producers such as seagrasses and macroalgae, detailed studies on the response of these benthic habitat components to drivers remain relatively sparse. Utilizing a multi-faceted examination of turtle-seagrass and sea urchin-macroalgae consumer and nutrient dynamics, I elucidate the impact of these drivers in Akumal, Quintana Roo, Mexico. In Yal Ku Lagoon, macroalgae bioindicators signified high nutrient availability, which is important for further studies, but did not consistently follow published trends reflecting decreased δ 15N content with distance from suspected source. In Akumal Bay, eutrophication and grazing by turtles and fishes combine to structure patches within the seagrass beds. Grazed seagrass patches had higher structural complexity and productivity than patches continually grazed by turtles and fishes. Results from this study indicate that patch abandonment may follow giving-up density theory, the first to be recorded in the marine environment. As Diadema antillarum populations recover after their massive mortality thirty years ago, the role these echinoids will have in reducing macroalgae cover and altering ecosystem state remains to be clear. Although Diadema antillarum densities within the coral reef ecosystem were comparable to other regions within the Caribbean, the echinoid population in Akumal Bay was an insufficient driver to prevent dominance of a turf-algal-sediment (TAS) state. After a four year study, declining coral cover coupled with increased algal cover suggests that the TAS-dominated state is likely to persist over time despite echinoid recovery. Studies on macroalgal diversity and nutrients within this same region of echinoids indicated diversity and nutrient content of macroalgae increased, which may further increase the persistence of the algal-dominated state. This study provides valuable insight into the variable effects of herbivores and nutrients on primary producers within a tropical coastal ecosystem. Results from this work challenge many of the currently accepted theories on primary producer response to nutrients and herbivory while providing a framework for further studies into these dynamics.

Top down control in a relatively pristine seagrass ecosystem

The loss of large-bodied herbivores and/or top predators has been associated with large-scale changes in terrestrial, freshwater, and marine ecosystems around the world. Understanding the consequences of these declines has been hampered by a lack of studies in relatively pristine systems. To fill this gap, I investigated the dynamics of the relatively pristine seagrass ecosystem of Shark Bay, Australia. I began by examining the seagrass species distributions, stoichiometry, and patterns of nutrient limitation across the whole of Shark Bay. Large areas were N-limited, P-limited, or limited by factors other than nutrients. Phosphorus-limitation was centered in areas of restricted water exchange with the ocean. Nutrient content of seagrasses varied seasonally, but the strength of seasonal responses were species-specific. Using a cafeteria-style experiment, I found that fast-growing seagrass species, which had higher nutrient content experienced higher rates of herbivory than slow-growing species that are dominant in the bay but have low nutrient content. Although removal rates correlated well with nutrient content at a broad scale, within fast-growing species removal rates were not closely tied to N or P content. Using a combination of stable isotope analysis and animal borne video, I found that green turtles (Chelonia mydas)—one of the most abundant large-bodied herbivores in Shark Bay—appear to assimilate little energy from seagrasses at the population level. There was, however, evidence of individual specialization in turtle diets with some individuals foraging largely on seagrasses and others feeding primarily on macroalgae and gelatinous macroplankton. Finally, I used exclusion cages, to examine whether predation-sensitive habitat shifts by megagrazers (green turtles, dugongs) transmitted a behavior-mediated trophic cascade (BMTC) between sharks and seagrasses. In general, data were consistent with predictions of a behavior-mediated trophic cascade. Megaherbivore impacts on seagrasses were large only in the microhabitat where megaherbivores congregate to reduce predation risk. My study highlights the importance of large herbivores in structuring seagrass communities and, more generally, suggests that roving top predators likely are important in structuring communities—and possibly ecosystems—through non-consumptive pathways.