The response of fishes to submerged aquatic vegetation complexity in two ecoregions of the mid-Atlantic bight: Buzzards Bay and Chesapeake Bay

Estuarine seagrass ecosystems provide important habitat for fish and invertebrates and changes in these systems may alter their ability to support fish. The response of fish assemblages to alteration of eelgrass (Zostera marina) ecosystems in two ecoregions of the Mid-Atlantic Bight (Buzzards Bay and Chesapeake Bay) was evaluated by sampling historical eelgrass sites that currently span a broad range of stress and habitat quality. In two widely separated ecoregions with very different fish faunas, degradation and loss of submerged aquatic vegetation (SAV) habitat has lead to declines in fish standing stock and species richness. The abundance, biomass, and species richness of the fish assemblage were significantly higher at sites that have high levels of eelgrass habitat complexity (biomass >100 wet g m^?2; density <100 shotts m^?2) compared to sites that have reduced eelgrass (biomass <100 wet g m^?2; density <100 shoots m^?2) or that have completely lost eelgrass. Abundance, biomass, and species richness at reduced eelgrass complexity sites also were more variable than at high eelgrass complexity habitats. Low SAV complexity sites had higher proportions of pelagic species that are not dependent on benthic habitat structure for feeding or refuge. Most species had greater abundance and were found more frequently at sites that have eelgrass. The replacement of SAV habitats by benthic macroalgae, which occurred in Buzzards Bay but not Chesapeake Bay, did not provide an equivalent habitat to seagrass. Nutrient enrichment-related degradation of eelgrass habitat has diminished the overall capacity of estuaries to support fish populations.     

Invasions in Marine Communities: Contrasting Species Richness and Community Composition Across Habitats and Salinity

While many studies of non-native species have examined either soft-bottom or hard-bottom marine communities, including artificial structures at docks and marinas, formal comparisons across these habitat types are rare. The number of non-indigenous species (NIS) may differ among habitats, due to differences in species delivery (trade history) and susceptibility to invasions. In this study, we quantitatively compared NIS to native species richness and distribution and examined community similarity across hard-bottom and soft-sediment habitats in San Francisco Bay, California (USA). Benthic invertebrates were sampled using settlement panels (hard-bottom habitats) and sediment grabs (soft-bottom habitats) in 13 paired sites, including eight in higher salinity areas and five in lower salinity areas during 2 years. Mean NIS richness was greatest in hard-bottom habitat at high salinity, being significantly higher than each (a) native species at high salinity and (b) NIS richness at low salinity. In contrast, mean NIS richness in soft-bottom communities was not significantly different from native species richness in either high- or low-salinity waters, nor was there a difference in NIS richness between salinities. For hard-bottom communities, NIS represented an average of 79% of total species richness per sample at high salinity and 78% at low salinity, whereas the comparable values for soft bottom were 46 and 60%, respectively. On average, NIS occurred at a significantly higher frequency (percent of samples) than native species for hard-bottom habitats at both salinities, but this was not the case for soft-bottom habitats. Finally, NIS contributed significantly to the existing community structure (dissimilarity) across habitat types and salinities. Our results show that NIS richness and occurrence frequency is highest in hard-bottom and high-salinity habitat for this Bay but also that NIS contribute strongly to species richness and community structure across each habitat evaluated.     

Environmental Determinants of Emergent Macrophyte Vegetation in Pacific Northwest Estuarine Tidal Wetlands

We investigated whether within wetland environmental conditions or surrounding land cover measured at multiple scales were more influential in structuring regional vegetation patterns in estuarine tidal wetlands in the Pacific Northwest, USA. Surrounding land cover was characterized at the 100, 250, and 1,000 m, and watershed buffer scales. Vegetation communities were characterized by high species richness, lack of monotypic zonation, and paucity of invasive species. The number of species per site ranged between 4 and 20 (mean?±?standard deviation?=?10.2?±?3.1). Sites supported a high richness (mean richness of native species 8.7?±?2.8) and abundance of native macrophytes (mean relative abundance 85 %?±?19 %). Vegetation assemblages were dominated by a mix of grasses, sedges, and herbs with Sarcocornia pacifica and Distichlis spicata being common at sites in the oceanic zone of the estuary and Carex lyngbyei and Agrostis stolonifera being common at the fresher sites throughout the study area. The vegetation community was most strongly correlated with salinity and land cover within close proximity to the study site and less so with land cover variables at the watershed scale. Total species richness and richness of native species were negatively correlated with the amount of wetland in the buffer at all scales, while abundance of invasive species was significantly correlated to within wetland factors, including salinity and dissolved phosphorus concentrations. Landscape factors related to anthropogenic disturbances were only important at the 100-m buffer scale, with anthropogenic disturbances further from the wetland not being influential in shaping the vegetation assemblage. Our research suggests that the traditional paradigms of tidal wetland vegetation structure and environmental determinants developed in east coast US tidal wetlands might not hold true for Pacific Northwest wetlands due to their unique chemical and physical factors, necessitating further detailed study of these systems.     

The effects of eelgrass habitat loss on estuarine fish communities of southern New England

We studied the late June–August fish community in extant and former eelgrass (Zostera marina L.) habitats in 15 estuaries of Buzzards Bay, and in Waquoit Bay, Massachusetts, U.S. Our objective was to quantify the effects of eelgrass habitat loss on fish abundance, biomass, species composition and richness, life-history characteristics, and habitat use by examining the response of the fish community to eelgrass loss in Waquoit and Buttermilk Bays over an 11-yr period (1988–1999) and in 14 other embayments of Buzzards Bay during 1993, 1996, and 1998. Sampling sites were located in present-day or historical eelgrass beds and were classified according to eelgrass habitat complexity (zero complexity: no eelgrass; low complexity: <100 eelgrass shoots or <100 g wet weight m⁻²; high complexity: ≥100 shoots and ≥100 g wet weight m⁻²). Habitats that had lost eelgrass included a variety of substratum types, from bare mud bottom to dense accumulations of red, brown, and green macroalgae (up to 7,065 g wet weight m⁻²). Contemporaneous sampling of fish (by otter trawl) and vegetated habitat (by divers) was conducted at each site. Overall, fish abundance, biomass, species richness, dominance, and life history diversity decreased significantly along the gradient of decreasing eelgrass habitat complexity. Loss of eelgrass was accompanied by significant declines in these measures of fish community integrity. Ten of the 13 most common species collected from 1988–1996 in Waquoit and Buttermilk Bays showed maximum abundance and biomass in sites with high eelgrass habitat complexity. All but two common species declined in abundance and biomass with the complete loss of eelgrass.     

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.     

Restored Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) as a Refuge for Epifaunal Biodiversity in Mid-Western Atlantic Coastal Bays

As nearshore ecosystems are increasingly degraded by human activities, active restoration is a critical strategy in ensuring the continued provision of goods and services by coastal habitats. After being absent for nearly six decades, over 1800 ha of the foundational species eelgrass (Zostera marina L.) has been successfully re-established in the coastal bays of the mid-western Atlantic, USA, but nothing is known about the recovery of associated animal communities in this region. Here, we determine the patterns and drivers of functional recovery in epifaunal invertebrates associated with the restored eelgrass habitat from 2001 to 2013. After less than a decade, the invertebrate community in the restored bed was richer, more even, and exhibited greater variation in functional traits than a nearby reference bed. Analysis of a suite of environmental and physical variables using random forests revealed these differences were primarily due to the increasing area and density of eelgrass, a direct consequence of ongoing restoration efforts. Based on analysis of functional traits, we propose that the rapid life histories of constituent organisms may have played a key role in their successful recovery. We also speculate that diverse epifaunal communities may have contributed to the restoration success through a well-described mutualism with eelgrass. Given that restored eelgrass now make up 32 % of total seagrass cover in the mid-Atlantic coastal bays, this restoration may conserve regional biodiversity by providing new and pristine habitat, particularly given the general decline of existing eelgrass in this region.     

Is Salinity Variability a Benthic Disturbance in Estuaries?

Freshwater inflow is a driver of the functioning of estuaries, and average salinity is usually measured to identify the effects of inflow in salinity-zone habitats. However, salinity variability could act as a disturbance by producing unstable habitats, leading to the question: is salinity variance an indicator of benthic disturbance, and therefore a driver of community stability? The macrofauna communities of five estuaries that lie in a climatic gradient on the Texas coastline were analyzed using a 26-year data set. Comparisons within and between estuaries with different inflow regimes were used as a natural experiment to simulate press disturbance events (i.e., climatic inflow) and pulse disturbance (i.e., floods) in maintaining community stability. Salinity average and variance was compared with benthic community diversity, evenness, and species richness. Salinity variance was more correlated to benthic diversity for each estuarine system (r?=??0.6610; p?=?0.0015) than average salinity (r?=?0.3818; p?=?0.0967). As salinity variance decreased (i.e., stability increased), diversity levels of benthic communities increased, and areas with mgore freshwater inflow displayed lower levels of benthic diversity. These findings advance a component of the general theory of diversity maintenance that persistent stressors, such as salinity variability, can influence diversity.     

Large-Scale Differences in Community Structure and Ecosystem Services of Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) Beds Across Three Regions in Eastern Canada

Eelgrass (Zostera marina) forms extensive beds in temperate coastal and estuarine environments worldwide and provides important ecosystem services, including habitat for a wide range of species as well as nutrient cycling and carbon storage. However, little is known about how eelgrass ecosystem structure and services differ naturally among regions. Using large-scale field surveys, we examined differences in eelgrass bed structure, carbon and nitrogen storage, community composition, and habitat services across three distinct regions in Eastern Canada. We focused on eelgrass beds with low anthropogenic impacts to compare natural differences. In addition, we analyzed the relationships of eelgrass bed structure with environmental conditions, and species composition with bed structure and environmental conditions, to elucidate potential drivers of observed differences. Our results indicate that regional differences in eelgrass bed structure were weakly correlated with water column properties, whereas differences in carbon and nitrogen storage were mainly driven by differences in eelgrass biomass. There were distinct regional differences in species composition and diversity, which were particularly linked to temperature, as well as eelgrass bed structure indicating differences in habitat provision. Our results highlight natural regional differences in ecosystem structure and services which could inform spatial management and conservation strategies for eelgrass beds.     

Composition,abundance, biomass,and production of macrofauna in a New England estuary: Comparisons among eelgrass meadows and other nursery habitats

Quantitative suction sampling was used to characterize and compare the species composition, abundance, biomass, and secondary production of macrofauna inhabiting intertidal mud-flat and sand-flat, eelgrass meadow, and salt-marsh-pool habitats in the Nauset Marsh complex, Cape Cod, Massachusetts (USA). Species richness and abundance were often greatest in eelgrass habitat, as was macroinvertebrate biomass and production. Most striking was the five to fifteen times greater rate of annual macrofaunal production in eelgrass habitat than elsewhere, with values ranging from approximately 23–139 g AFDW m² yr^?1. The marsh pool containing widgeon grass (Ruppia maritima) supported surprisingly low numbers of macroinvertebrates, probably due to stressfully low dissolved oxygen levels at night during the summer. Two species of macroinvertebrates, blue mussels (Mytilus edulis) and to a lesser extent bay scallops (Argopecten irradians), used eelgrass as “nursery habitat.” Calculations showed that macroinvertebrate production is proportionally much greater than the amount of primary production attributable to eelgrass in the Nauset Marsh system, and that dramatic changes at all trophic levels could be expected if large changes in seagrass abundance should occur. This work further underscores the extraordinarily large impact that seagrass can have on both the structure and function of estuarine ecosystems. *** DIRECT SUPPORT *** A01BY070 00006     

Patterns of seasonal availability and habitat use by fishes and decapod crustaceans in a southern New Jersey estuary

We examined the community structure of fish and selected decapod crustaceans and tested for within estuary differences among habitats at depths of 0.6 m to 7.9 m, in Great Bay and Little Egg Harbor in southern New Jersey. Several habitat types were identified a priori (e.g., eelgrass, sea lettuce, and marsh creeks) and sampled by trawl (4.9 m headrope, 19-mm mesh wings, 6.3-mm mesh liner), monthly, from June 1988 through October 1989. Repetitive (n=4) 2-min trawl tows were taken at each habitat type from 13 locations. The fishes and decapod crustaceans collected were typical of other Mid-Atlantic Bight estuaries but varied greatly inseasonal abundance and species. In the years sampled, bay anchovy (Anchoa mitchilli) was the dominant species (50.5% of the total number), followed by spot (Leiostomus xanthurus) (10.7%), Atlantic silverside (Menidia menidia) (9.7%), fourspine stickleback (Apeltes quadracus) (5.9%), blue crab (Callinectes sapidus) (4.6%), and northern pipefish (Syngnathus fuscus) (4.2%). The biota were examined by multi-dimensional scaling (MDS) for habitat associations and “best abiotic predictor” of community structure. Percent silt combined with salinity was the most important abiotic determinant of the faunal distributions among habitats. Temperature was a major factor influencing seasonal occurrence of the biota but had less effect on habitat comparisons. The analysis confirmed the distinct nature of the assemblages associated with the habitats, that is, eelgrass, upper estuary subtidal creeks, channels, and open bay areas. Several species were associated with specific habitats: for example,A. quadracus andS. fuscus with eelgrass, clupeids with subtidal creek stations,L. xanthurus with marsh channels, and black sea bass (Centropristis striata) and spotted hake (Urophycis regia) with sponge-peat habitat. Species richness appeared to be positively related to habitat structural heterogeneity. Thus, the best predictors for these estuarine fish and decapod crustacean assemblages were seasonal temperature, percent silt and salinity combined, and the physical heterogeneity of the habitat.     

Structural components of eelgrass (Zostera marina) meadows in the lower Chesapeake Bay—Decapod crustacea

Otter trawl collections of eelgrass habitats in the lower Chesapeake Bay during 1976–1977 produced 14 species of decapod crustaceans. These collections were dominated by palaemonid shrimp (Palaemonetes spp.), blue crabs (Callinectes sapidus), and sand shrimp (Crangon septemspinosa), each of which exhibited unimodal seasonal abundance curves with large summer peaks. Decapod abundance was positively correlated with plant biomass throughout the year. Decapod densities on vegetated bottoms were greater than on unvegetated bottoms, and nighttime abundance on each bottom type was greater than corresponding daytime abundance. Total decapod abundances in Chesapeake Bay eelgrass meadows appear to be much greater than those reported in North Carolina eelgrass or Gulf of Mexico turtlegrass habitats.     

Epibenthic fishes and decapod crustaceans in northern estuaries: A comparison of vegetated and unvegetated habitats in maine

Species richness and abundance of epibenthic fishes and decapod crustaceans were quantified with day-time beam trawl tows and throw traps to provide information on nekton assemblages inZostera marina and unvegetated sandy habitats in northern latitudes. Sampling at randomly selected stations with a 1.0-m beam trawl occurred in eelgrass (Zostera marina) and unvegetated sandy substrates of two mid-coastal Maine estuaries: Casco Bay and Weskeag River. Random 1.0-m throw trap samples were collected inZostera and adjacent unvegetated sandy substrates in Casco Bay and Weskeag River as well. Species richness and faunal abundances were positively associated with the occurrence ofZostera within Weskeag River and Casco Bay estuaries using both gear types. A total of 17 species of fishes and 6 species of decapods were collected in the two estuaries using both gears. Populations of most species were dominated by young-of-the-year and juvenile life history stages. Number and densities of fishes were higher inZostera, due primarily to the abundances of eelgrass residents such as threespine,Gasterosteus aculeatus, and fourspine sticklebacks,Apeltes quadracus, grubby,Myoxocephalus aenaeus, and cunner,Tautogolabrus adspersus. Crangon septemspinosa dominated decapod catch per unit effort and density in both estuaries and habitats.     

The effect of aquaculture practices on the benthic macroinvertebrate community of a lagoon system in the Bay of Cádiz (southwestern Spain)

Monthly quantitative Ekman-Birge grab sampling was used to characterize and compare the composition and structure of the benthic macroinvertebrate community inhabiting semi-enclosed polyculture lagoons (SPL) (three sampling sites) and enclosed monoculture ponds (EMP) (two sampling sites) of a lagoonal system of the Bay of Cádiz. The two areas differed considerably in habitat characteristics and aquaculture management. The SPL area was characterized by low rates of water exchange, low fish densities, and the presence of a macroalgal cover. In the EMP area, there was a complete exchange of water daily (by pumping) and a supply of food pellets, density of fish was high, and no vegetative cover was present. There were considerable differences in species composition between habitats with different culture methods: 11 of the 21 most abundant species were exclusive to one or the other. Several epibenthic species were abundant in the polyculture lagoon but were low in density or were absent in monoculture ponds. Some infaunal species, on the other hand, were more abundant in the monoculture ponds. Univeriate measures of community structure (abundance and biomass, Margalef’s species richness, Shannon-Wiener diversity, and Pielou’s evenness indices) did not indicate significant differences between the SPL and EMP areas. Conversely, the abundance-biomass comparison (ABC) method indicated that, on average, the macrobenthic community was moderately disturbed in the SPL and undisturbed in the EMP areas. Multidimensional scaling (MDS) ordination and hierarchical cluster analysis (Bray-Curtis similarity measure) revealed the occurrence of two main benthic assemblages that corresponded to the aquaculture methods. The different rates of water exchange for the two aquaculture practices seem to have contributed to differences in the composition and structure of the benthic communities.     

An estuarine benthic index of biotic integrity (B-IBI) for Chesapeake Bay

A multimetric benthic index of biotic integrity (B-IBI) was developed using data from five Chesapeake Bay sampling programs conducted between 1972 and 1991. Attributes of the index were selected by comparing the response of 17 candidate measures of benthic condition (metrics) between a set of minimally affected reference sites and at all other sites for which data were available. This procedure was conducted independently for each of seven habitats defined by salinity and substrate. Fifteen of the 17 candidate metrics differed significantly between reference sites and other sites for at least one habitat. No metric differed significantly in all seven habitats; however, four metrics, species diversity, abundance, biomass, and percent of abundance as pollution-indicative taxa, differed in six habitats. The index was calculated by scoring each selected metric as 5, 3, or 1 depending on whether its value at a site approximated, deviated slightly from, or deviated greatly from conditions at the best reference sites. Validation based on independent data collected between 1992 and 1994 indicated that the index correctly distinguished stressed sites from reference sites 93% of the time, with the highest validation rates occurring in high salinity habitats.     

Fishes and decapod crustaceans of Cape Cod eelgrass meadows: Species composition,seasonal abundance patterns and comparison with unvegetated substrates

Bimonthly trawl samples from eelgrass and nearby unvegetated areas on Cape Cod, Massachusetts, showed greater species richness in eelgrass meadows relative to unvegetated areas, and greater summer abundance in vegetation for decapod crustaceans and fishes. The composition of eelgrass-associated decapods and fishes was dominated by cold-water taxa and was strikingly different from that of the better studied eelgrass meadows of the mid-Atlantic coast. Four of the eight decapod species collected, including the second and third most abundant taxa, do not even appear in collections reported from Chesapeake Bay eelgrass meadows. Similarly, 10 of the 22 fish species taken, including the first and sixth most abundant species, are not reported from Chesapeake Bay eelgrass samples. Cape Cod eelgrass beds seem to play a nursery role for several commercially important fish species, although the nursery function is less obvious than in previously studied mid-Atlantic eelgrass meadows.     

Expression of the Estuarine species minimum in littoral fish assemblages of the Lower Chesapeake Bay Tributaries

Species richness declines to a minimum (artenminimum) in the oligohaline reach of estuaries and other large bodies of brackish water. To date, observations of this feature in temperate estuaries have been largely restricted to benthic macroinvertebrates. Five years of seine data collected during the summers of 1990–1995 in the major tidal tributaries to the lower Chesapeake Bay were examined to see if this feature arose in estuarine fish assemblages. Estimates of numerical species richness (alpha diversity) and rates of species turnover between sites (beta diversity) were generated via rarefaction and detrended correspondence analysis. Two spatial attributes of the distribution of littoral fish species along salinity gradients in the tributaries of the lower Chesapeake Bay were revealed: (1) a species richness depression in salinities of 8–10% and (2) a peak in the rate of species turnover associated with the tidal freshwater interface (salinities of 0–2%). Expression of the minimum is influenced by the physical length of the salinity gradient and the interaction between a species’ salinity preferences and tendency to make long excursions from favorable habitats.     

Diverse Dietary Responses by Saltmarsh Consumers to Chronic Nutrient Enrichment

We examined the effect of whole-ecosystem nutrient enrichment on herbivory in saltmarsh creek-wall habitats in the Plum Island Estuary (Massachusetts, USA). Located between the macrophyte-dominated high marsh and adjoining mudflats, creek walls are steep vertical habitats vegetated with productive filamentous algae and associated epiphytes. Annual nitrate and phosphate loading rates were increased approximately ×10–15 in creeks mimicking short-term (2-month) and chronic (6-year) eutrophication. We assessed the diets of epifaunal invertebrates (three gastropods and one amphipod species) that potentially graze on benthic algae using natural isotope abundance data and per capita grazing rate measurements derived from ¹³C prelabeled algae. Substantial dietary contributions from benthic algae were observed in all consumers even though previous research has indicated most rely on Spartina detritus as the principal food resource. The amphipod Orchestia grillus and the snail Melampus bidentatus grazed benthic algae in excess of 500 μg algal C g C^?1 h^?1, whereas the snail Nassarius obsoletus and hydrobiid snails grazed at lower rates. Few dietary changes were detected with short-term enrichment. Algal grazing rates of N. obsoletus and M. bidentatus increased with chronic enrichment probably as a functional response to increased algal productivity. O. grillus grazed at a high rate and parasitic infection did not affect its consumption of benthic algae. The abundance and frequency of occurrence of O. grillus on creek-wall habitats increased with chronic nutrient enrichment suggesting amphipods contribute to top–down control on benthic algae and slow algal growth as nutrient enrichment occurs.     

Structural components of eelgrass (Zostera marina) meadows in the lower Chesapeake Bay—Fishes

The structure of the fish community associated with eelgrass beds in the lower Chesapeake Bay was studied over a 14 month period. A total of 24,182 individuals in 48 species was collected by otter trawl with Leiostomus xanthurus (spot) comprising 63% of the collection, Syngnathus fuscus (northern pipefish) 14%, Anchoa mitchilli (bay anchovy) 9%, and Bairdiella chrysoura (silver perch) 5%. The density and diversity of fishes were higher in vegetated areas compared to unvegetated areas; fishes were more abundant in night collections Fish abundance and species number increased in the spring and early summer as both water temperature and eelgrass biomass increased and decreased in the fall and winter as temperature and eelgrass biomass decreased. Gill netting revealed some of the top predators in the system, especially the sandbar shark, Carcharhinus milberti. The fish community in the Chesapeake Bay was quite different from North Carolina eelgrass fish communities. Most notable was the rarity of the pinfish, Lagodon rhomboides, which may be a very important predator in the structuring of the epifaunal communities.     

Factors influencing spatial and annual variability in eelgrass (Zostera marina L.) meadows in Willapa Bay, Washington, and Coos Bay, Oregon, estuaries

Environmental factors that influence annual variability and spatial differences (within and between estuaries) in eelgrass meadows (Zostera marine L.) were examined within Willapa Bay, Washington, and Coos Bay, Oregon, over a period of 4 years (1998–2001). A suite of eelgrass metrics were recorded annually at field sites that spanned the estuarine gradient from the marine-dominated to mesohaline region of each estuary. Plant density (shoots m^?2) of eelgrass was positively correlated with summer estuarine salinity and inversely correlated with water temperature gradients in the estuaries. Eelgrass density, biomass, and the incidence of flowering plants all increased substantially in Willapa Bay, and less so in Coos Bay, over the duration of the study. Warmer winters and cooler summers associated with the transition from El Niño to La Niña ocean conditions during the study period corresponded with this increase in eelgrass abundance and flowering. Large-scale changes in climate and nearshore ocean conditions may exert a strong regional influence on eelgrass abundance that can vary annually by as much as 700% in Willapa Bay. Lower levels of annual variability observed in Coos Bay may be due to the stronger and more direct influence of the nearshore Pacific Ocean on the Coos Bay study sites. The results suggest profound effects of climate variation on the abundance and flowering of eelgrass in Pacific Northwest coastal estuaries.     

Macroalgal distribution patterns in a shallow, soft-bottom lagoon, with emphasis on the nonnativeGracilaria vermiculophylla andCodium fragile

We determined the distribution of macroalgae in Hog Island Bay, a shallow coastal lagoon in Virginia, USA, seasonally at 12 sites from 1998 to 2000 and at 3 representative sites from 2000 to 2002. We analyzed macroalgal biomass, taxonomic richness, and abundance of two non-native species, the cryptic invaderGracilaria vermiculophylla and the conspicuousCodium fragile, with respect to season, location (mainland, mid lagoon, barrier island sites), and elevation (intertidal, subtidal). Taxonomic richness, total algal biomass, and nonnative biomass peaked in the summer months when temperature and light availability were highest. A few stress tolerant and ephemeral algae dominated the algal assemblage.G. vermiculophylla constituted 74% of the entire algal biomass, was the most abundant alga in all seasons, locations, and elevation levels, and was positively correlated with taxonomic richness and abundance of filamentous species.Ulva curvata, Bryopsis plumosa, andC. fragile accounted for an additional 16% of the algal biomass. There are distinct habitats in Hog Island Bay that can be classified into low diversity-low biomass regions near the mainland and barrier islands and high diversity-high biomass regions in the open mid lagoon, where abundant shells for attachment and intermediate levels of water column nutrients and turbidity likely create better growth conditions. Taxonomic richness and biomass were higher in subtidal than intertidal zones, presumably due to lower desiccation stress. This study provides an example of how a single invasive species can dominate an entire assemblage, both in terms of biomass (being most abundant in all seasons, locations, and tidal levels) and species richness (correlating positively with epiphytic filamentous taxa). By adding hard-substratum structural complexity to a relatively homogenous soft-substratum system,G. vermiculophylla increases substratum availability for attachment and entanglement of other algal species and enhances local diversity. Without widespread and abundantG. vermiculophylla, taxa likePolysiphonia, Ceramium, Bryopsis, Ectocarpus, andChampia would likely be much less common. This study also highlights the importance of using DNA analysis of voucher specimens in monitoring programs to accurately identify cryptic invaders.