Organic carbon metabolism and carbonate dynamics in a Mediterranean seagrass (Posidonia oceanica), meadow

We measured monthly dissolved oxygen (DO) changes in situ benthic incubations from March 2001 to October 2002 in aPosidonia oceanica meadow and unvegetated sediments of Magalluf Bay (Mallorca Island, Spain) to determine gross primary production (GPP), community respiration (R), and net community production (NCP). From June 2001 to October 2002, we also measured fluxes of dissolved inorganic carbon (DIC) and total alkalinity (TAlk). The yearly integrated metabolic rates based on DO changes show that theP. oceanica communities are net autotrophic while the metabolic rates in the unvegetated benthic communities are nearly balanced. Higher calcium carbonate (CaCO₃) cycling, both in terms of production and dissolution, was observed inP. oceanica communities than in unvegetated benthic communities. In theP. oceanica meadow, the annual release of CO₂ from net CaCO₃ production corresponds to almost half of the CO₂ uptake by NCP based on DIC incubations. In unvegetated benthic communities, the annual uptake of CO₂ from net CaCO₃ dissolution almost fully compensates the CO₂ release by NCP based on DIC incubations. CaCO₃ dynamics is potentially a major factor in CO₂ benthic fluxes in seagrass and carbonate-rich temperate coastal ecosystems.     

Direct evidence of imbalanced seagrass (Posidonia oceanica) shoot population dynamics in the Spanish Mediterranean

Direct census of shoots tagged in permanent plots was used to assess the present (2000–2002)Posidonia oceanica population dynamics in 25 meadows along the Spanish Mediterranean Coast. Shoot density ranged from 154±8 to 1,551±454 shoots m⁻², absolute shoot mortality from 5±0 to 249±53 shoots m⁻² yr⁻¹, and absolute shoot recruitment from <5 ±1 to 62±42 shoots m⁻²yr⁻¹. Specific shoot mortality and recruitment rates, which are mathematically and statistically (p>0.05) independent of shoot density, varied from 0.015±0.006 to 0.282±0.138 yr⁻¹ and 0.018±0.005 to 0.302±0.093 yr⁻¹, respectively. Absolute shoot mortality rate was scaled to shoot density (Pearson correlation, r=0.78, p<0.0001), and variability in specific shoot recruitment rate was partially due to differences in the percentage of growing apexes, which produce most of the recruits within the population (Pearson correlation, r=0.50, p<0.001), demonstrating the existence of structural constraints on shoot demography. Shoot half-life was estimated to range from 2.5 to 60.4 yr and meadow turnover times between 6.7 yr and more than a century, provided current estimates of shoot mortality, recruitment rates, and density remain uniform. There were differences in shoot mortality and recruitment at the regional scale, with the meadows developing along the coast of the Spanish mainland experiencing the highest shoot mortality (Tukey test, p<0.05) and tending to exhibit the highest shoot recruitment. The low shoot recruitment did not balance shoot mortality in most (60%) of the meadows, showing a prevalence of declining populations among the 25 meadows studied (Wilcoxon ranked sign test, p<0.0005). This study demonstrates the power of direct census of seagrass shoots in permanent plots to evaluate the present status of seagrass meadows, to detect on-going population decline, and to provide some insight onto the possible factors involved. The incorporation of direct census of seagrass meadows to monitoring programs will help provide the early-warning signals necessary to support management decisions to conserve seagrass meadows.     

Epiphyte-grazer relationships in seagrass meadows: Consequences for seagrass growth and production

Studies of seagrass meadows have shown that the production of algal epiphytes attached to seagrass blades approaches 20% of the seagrass production and that epiphytes are more important as food for associated fauna than are the more refractory seagrass blades. Since epiphytes may compete with seagrasses for light and water column nutrients, excessive epiphytic fouling could have serious consequences for seagrass growth. We summarize much of the literature on epiphytegrazer relationships in seagrass meadows within the context of seagrass growth and production. We also provide insights from mathematical modeling simulations of these relationships for a Chesapeake BayZostera marina meadow. Finally we focus on future research needs for more completely understanding the influences that epiphyte grazers have on seagrass production.     

Comparison of restored and natural seagrass beds near Corpus Christi,Texas

Structural equivalence between seagrass restoration sites and adjacent natural seagrass beds on the mid Texas coast was assessed six times between April 1995 and May 1997. Throw traps and corers were used for quantitative sampling. Restoration sites were 2.7 to 6.6 yr old when first sampled and 3.7 to 8.2 yr old when last sampled. There were few significant differences in water column, seagrass, or sediment characteristics, in fish and decapod (nekton) densities, or in nekton and benthos community compositions between restored and natural seagrass habitats at any time during the study period. Differences in densities of dominant benthic invertebrates were regularly observed, with greater densities of more taxa observed in natural seagrasses than in restored beds. Densities of Class Oligochaeta and the polychaetePrionospio heterobranchiata are proposed as potential indicators of structural equivalence in restored seagrasses. This study indicates that seagrass restorations in the vicinity of Corpus Christi, Texas, exhibit minimal quantitative differences in community structure (except for benthos) relative to adjacent natural seagrass beds after 3 to 5 yr.     

Biogeochemical consequences of macrofauna burrow ventilation

The burrow walls created by macrofauna in aquatic sediments are sites of intense chemical mass transfer. Quantitative measurement of their significance is, however, difficult because chemistry in the immediate vicinity of burrow walls is temporally dynamic due to periodic ventilation of burrows by macrofauna. A temporally dynamic, 2D multicomponent diffusion-reaction model was utilized to depict the magnitude and time dependency of chemical mass transfer in the immediate vicinity of burrow walls as well as at the water/sediment interface. The simulation results illustrate that sediment particles, pore water, and microorganisms within a few millimeters of burrow walls experience significant oscillation in pH (as much as two pH units) and dissolved oxygen concentration (between saturation and near anoxia) whereas such oscillation is absent at the water/ sediment interface. The geochemical oscillation is expected to affect the net stability of mineral phases, activities and community structures of microorganisms, and rates and magnitudes of microbial diagenetic reactions.     

Comparison of stream sediment and soil sampling for regional exploration in the eastern Alps,Austria

Tests in three different areas of the Austrian Alps showed that in mountainous areas, soil samples taken at the break of slope are an alternative sampling medium to stream sediments for regional geochemical exploration even if taken at the unusually low density of 1 sample/km². B-horizon soil samples from the nearest break of slope were collected in conjunction with stream sediment orientation studies. The minus 0.2 mm fraction of all samples was routinely analyzed for the total contents of 36 elements. The results for the indicator elements in the three test areas (Area 1: Cu; Area 2: Ag, As, Cu, Pb, Sb, Sn, Zn; Area 3: As, Cu) showed that break-of-slope soil samples more clearly indicate known mineralization than drainage geochemistry. In the soils, almost all elements showed a higher natural variability of the data with considerably higher metal contents. In mountaineous regions, each soil sample will have a clearly defined area of influence. The break of slope represents a site where clastically and hydromorphically dispersed elements are concentrated. It is concluded that soil samples can be used for regional exploration using low sampling densities in mountaineous areas. At the same time they will provide valuable information for other fields of science (e.g. agriculture, forestry, environmental sciences) and are thus superior to stream sediments.     

Modeling nonlinear seagrass clonal growth: Assessing the efficiency of space occupation across the seagrass flora

The clonal growth of 9 seagrass species was modeled using a simulation model based on observed clonal growth rules (i.e., spacer length, rhizome elongation rates, branching rates, branching angle) and shoot mortality rates for seagrass species. The results of the model confirmed the occurrence of complex, nonlinear growth of seagrass clones derived from internal dynamics of space occupation. The modeled clones progressed from a diffuse-limited aggregation (DLA), dendritic growth, identified with a guerrilla strategy of space occupation, to a compact (Eden) growth, comparable to the phalanx strategy of space occupation, once internal recolonization of gaps, left by dead shoots within the clone, begins. The time at which seagrass clones shifted from diffuse limited to compact growth was predictable from the branching angle and frequency of the species and varied from 1 yr to several decades among species. As a consequence the growth behavior and the apparent growth strategy of the species changes with the development of the clones. The results of the model demonstrate that the emergent complexity of seagrass clonal growth is contained within the simple set of growth rules that can be used to represent clonal growth.     

Role of larger herbvores in seagrass communities

The nutritional ecology of macroherbivores in seagrass meadows and the roles of grazing by urchins, fishes and green turtles in tropical systems and waterfowl in temperate systems are discussed in this review. Only a few species of animals graze on living seagrasses, and apparently only a small portion of the energy and nutrients in seagrasses is usually channeled through these herbivores. The general paucity of direct seagrass grazers may be a function of several factors in the composition of seagrasses, including availability of nitrogen compounds, presence of relatively high amounts of structural cell walls, and presence of toxic or inhibitory substances. The macroherbivores, however, can have a profound effect on the seagrass plants, on other grazers and fauna associated with the meadow, and on chemical and decompositional processes occurring within the meadow. Grazing can alter the nutrient content and digestibility of the plant, as well as its productivity. Removal of leaf material can influence interrelations among permanent and transient faunal residents. Grazing also interrupts the detritus cycle. Possible consequences of this disruption, either through acceleration or through decreased source input, and the enhancement of intersystem coupling by increased export and offsite fecal production, are discussed. The extent and magnitude of these effects and their ecological significance in the overall functioning of seagrass meadows only can be speculated, and probably are not uniform or of similar importance in both tropical and temperate seagrass systems. However, areas grazed by large herbivores provide natural experiments in which to test hypotheses on many functional relations in seagrass meadows.     

Impacts of seagrass habitat architecture on bivalve settlement

We investigated the effects of differing spatial scales of seagrass habitat architecture on the composition and abundance of settling bivalves in a sub-tropical seagrass community. The density of newly settled bivalves was generally greater atThalassia testudinum grass bed edge (<1 m) compared to interior portions of the bed (>10 m). Deviation from this generalized pattern occurred when high densities of newly settled tulip mussels (Modiolus americanus) were recorded from the interior of the meadow, associated with aggregations of adult mussels. Bivalve settling densities appear to reflect settlement shadows of passively delivered larvae, bedload transport of newly settled individuals from unvegetated regions, as well as gregarious settlement among adult conspecifics. We also investigated the impact of seagrass patch shape and size on settlement by using artificial seagrass units (ASU) in separate short-term and long-term experiments. We found a positive relationship between ASU perimeter and bivalve abundance, suggesting that larval encounter rates with seagrass habitat may determine initial settlement patterns. Using ASUs we also investigated the relative role seagrass epiphytes play in determining the density of settling bivalves. Results showed greater settling densities where epiphytic secondary structure was elevated compared to controls, and bivalve density was significantly greater when ASUs were fouled with a natural community of epiphytes, suggesting that both microstructure and biofilms positively influenced bivalve settlement. We conclude that structural components of seagrass habitats increase bivalve settlement at multiple spatial scales, including epiphytic micro-structure, small-scale patch shape and size, and large-scale within habitat differences.     

The effects of tributyltin within aThalassia seagrass ecosystem

Flow-through seagrass core microcosms were used to examine responses of species and processes to a logarithmic gradient of dosing with¹⁴C-labeled tributyltin-chloride (TBT-CI). Experiments involved delivery of TBT-CI to the water column of replicate cores of a treatment (n=16) once per week; one-half of the cores were sacrificed after 3 wk of dosing, the others were dosed for 6 wk. Initial water column concentrations for the three treatments averaged 0.205, 2.23, and 22.21 μg I^?1, expressed as the TBT⁺ cation, but these concentrations dropped rapidly. Retained¹⁴C tracer, an estimate of total organotin species, was distributed to sediments, plants, and other biological tissues, all of whose tracer concentrations increased with time. Measures to indicate responses of both autotrophic and heterotrophic organisms were made; in general, treatment effects were demonstrable statistically only at the highest dose level. Accumulation of chlorophyll and biomass on glass slides was highest when suspended for the entire experiment in the water of the highest treatment; this unexpected result was perhaps an indirect effect related to reduced grazing activity in the microcosms. The highest dose of TBT-CI resulted in virtual population mortality of a few macrobenthic species and decreased loss of plant material in litter bags, both demonstrated within the first 3 wk of dosing. Reduced litter loss was coincident with mortality of an amphipod (Cymadusa compta) capable of shredding plant material, and a causal relation between the two effects is plausible. Thus, if concentrated to similar levels in aThalassia bed, TBT⁺ may have direct species-level effects and process-level effects, potentially causing ecosystem change via disruption of a species-process linkage influential in seagrass detrital food web dynamics.     

Setting load limits for nutrients and suspended solids based upon seagrass depth-limit targets

Total nitrogen (TN), total phosphorus (TP), and total suspended solids (TSS) loadings [log (kg ha⁻¹ yr⁻¹)] were regressed against seagrass depth limits (percent of depth-limit targets) to back-predict the load limits or allocations (kg ha⁻¹ yr⁻¹ or kg yr⁻¹) necessary to meet targeted seagrass depth limits in the Indian River and Banana River (IRBR) lagoons, Florida. Because the load allocations can be applied as total maximum daily loads (TMDL) for the IRBR (U.S. Environmental Protection Agency mandate), the method and results are developed and presented toward that end. The regression analyses indicate that the range of surface-discharge load limits (nonpoint + point source), per watershed area, required to achieve the desired depth limits for seagrass in the IRBR are approximately 2.4–3.2 kg ha⁻¹ yr⁻¹ TN, 0.41–0.64 kg ha⁻¹ yr⁻¹ TP, and 48–64 kg ha⁻¹ yr⁻¹ TSS. This simple regression method may have application to other shallow estuarine lagoons or bays where seagrass growth is limited by light and water transparency, water transparency is strongly affected by watershed pollutant loadings, water residence times are sufficiently long to allow seagrass coverage to respond to and covary with total load inputs, and multiyear monitoring has yielded sufficient variability in both pollutant loadings and seagrass coverages to develop a statistically meaningful relationship.     

A reassessment of the status of the benthic macrofauna of the Raritan estuary

The benthic macrofauna of the Raritan estuary, at the mouth of New York Harbor, has been reported to be severely impacted by pollution, with a significant change occurring between surveys in the late 1950’s and the early 1970’s. We review this assessment using five-fold more macrofaunal data (including biomass), not reported or used before, from a 1973–1974 survey. These data are compared to previous and similar studies in the area. This reassessment suggests benthic community structure is not as degraded as previously reported and is, in many ways, similar to other relatively unpolluted middle Atlanic estuaries.     

Testing the predictive power of seagrass depth limit models

The power of equations predicting seagrass depth limit (Z_c) from light extinction (K _z) was tested on data on seagrass depth limits collected from the literature. The test data set comprised 424 reports of seagrass colonization depth and water transparency, including data for 10 seagrass species. This data set confirmed the strong negative relationship betweenZ _c andK _z. The regression equation in Duarte (1991) overestimated the realized seagrass colonization depths at colonization depths < 5 m, while there was no prediction bias above this threshold. These results indicated that seagrass colonizing turbid waters (K _z 0.27 m^-1) have higher apparent light requirements than those growing in clearer waters. The relationship between seagrass colonization depth and light attenuation shifts at a threshold of light attenuation of 0.27 m^-1, requiring separate equations to predictZ _c for seagrass growing in more turbid waters and clearer waters, and to set targets for seagrass restoration and conservation efforts.     

Patterns of seagrass community response to local shoreline development

Three quarters of the global human population will live in coastal areas in the coming decades and will continue to develop these areas as population density increases. Anthropogenic stressors from this coastal development may lead to fragmented habitats, altered food webs, changes in sediment characteristics, and loss of near-shore vegetated habitats. Seagrass systems are important vegetated estuarine habitats that are vulnerable to anthropogenic stressors, but provide valuable ecosystem functions. Key to maintaining these habitats that filter water, stabilize sediments, and provide refuge to juvenile animals is an understanding of the impacts of local coastal development. To assess development impacts in seagrass communities, we surveyed 20 seagrass beds in lower Chesapeake Bay, VA. We sampled primary producers, consumers, water quality, and sediment characteristics in seagrass beds, and characterized development along the adjacent shoreline using land cover data. Overall, we could not detect effects of local coastal development on these seagrass communities. Seagrass biomass varied only between sites, and was positively correlated with sediment organic matter. Epiphytic algal biomass and epibiont (epifauna and epiphyte) community composition varied between western and eastern regions of the bay. But, neither eelgrass (Zostera marina) leaf nitrogen (a proxy for integrated nitrogen loading), crustacean grazer biomass, epifaunal predator abundance, nor fish and crab abundance differed significantly among sites or regions. Overall, factors operating on different scales appear to drive primary producers, seagrass-associated faunal communities, and sediment properties in these important submerged vegetated habitats in lower Chesapeake Bay.     

The Holocene fossil marine macrofauna of the Vestfold Hills, East Antarctica

The Holocene (< 10,000 years old) fossil marine macrofauna (molluscs, polychaetes, mammals and birds) of the Vestfold Hills, Antarctica (68°30'S, 78°00'E) is described. All species are extant, with circumantarctic or wider distributions and wide depth ranges. At Heidemann Bay, Pleistocene-age (> 30 ky old) shell fragments occur in a Holocene moraine. These are interpreted as being glacially reworked fossil shells. Holocene benthic estuarine and coastal molluses and serpulid worm tubes are abundant on the flanks of terraces above saline and hypersaline lakes. Extensive radiocarbon dating gives a chronology of these deposits which were not synchronous. Mummies of seals and penguins, which are concentrated on the shores of the saline lakes and scattered across the Hills, are of no use in dating geomorphological events. Marine fossils in moraines are difficult to interpret because they can be glacially-transported live or fossil shells, or deposited after wind transport. The fossil fauna has such a wide ecological range that it indicates only in general terms that for part of the Pleistocene and most of the Holocene, climatic conditions were broadly similar to the present.     

Experimental studies of predation on polychaetes associated with seagrass beds

Predation on benthic polychaetes associated with seagrass beds was examined in laboratory and field predator inclusion experiments with one fish and three invertebrate predators. Predation had differential effects on polychaete taxa which depended on their microhabitat utilization patterns. The magnitude of predation effects strongly depended on the predator species, with the shrimpPenaeus duorarum having the greatest impact and the pinfishLagodon rhomboides the least. Abundance of spionid polychaetes, which are located near the sediment-water interface, was reduced most often by predation. Nereid and capitellid polychaete abundances were reduced by some predators and not by others. The results emphasize the need for more detailed natural history information on polychaete species in order to improve the interpretation of results from predation experiments.     

Evidence of direct particle trapping by a tropical seagrass meadow

The capacity of seagrass canopies to directly retain sestonic particles was tested by quantifying the rate at which suspended fluorescent tracer particles were retained within a tropical Philippine seagrass meadow and by examining whether the test particles lost from the water column were later bound to seagrass leaves or inside epibionts. The particle loss rates in the presence of seagrass canopies were up to 4 times higher than those in unvegetated and plankton controls. The seagrass canopies trapped particles with a maximum rate of 0.73 (±0.24) h^?1. As much as 5% of the particles trapped by the seagrass leaves were physically adhered to the leaf surfaces following rinsing. Particles were also observed to be ingested by protozoa (ciliates and amoeba-like organisms), residing on the surface of the leaves, and may be the dominant particle trapping mechanism by seagrass leaves. These processes should provide an efficient mechanism for the transfer of planktonic production to the benthos, adding to the high organic carbon input maintained by the high production of the seagrass themselves.     

Some exact sampling distributions for variogram estimators

For equally spaced observations from a one-dimensional, stationary, Gaussian random function, the characteristic function of the usual variogram estimator for a fixed lag k is derived. Because the characteristic function and the probability density function form a Fourier integral pair, it is possible to tabulate the sampling distribution of a function of a using either analytic or numerical methods. An example of one such tabulation is given for an underlying model that is simple transitive.