Bacterial abundance and aerobic microbial activity across natural and oyster aquaculture habitats during summer conditions in a northeastern Pacific estuary

We measured sediment properties and the abundance and aerobic metabolism of microbes in Willapa Bay, Washington, USA, to test the response of sediment microbes to oyster aquaculture. Sites spanned the estuary gradient (practical salinity units ranged from 24 to 30 under seasonally low river flows) and six different low-intertidal habitat types: eelgrass (Zostera marina), unstructured tideflat, oyster hummocks (reefs of Crassostrea gigas), longline oyster aquaculture, hand-picked on-bottom oyster aquaculture, and dredged on-bottom oyster aquaculture. Aerobic metabolism was assessed by sole-source carbon use (SSCU) of 31 carbon sources on Biolog plates. Sediments generally became siltier and more organically enriched into the estuary, but no consistent differences in sediment properties occurred across habitat types. Bacterial cell density tracked organic content. Across the estuary gradient, overall aerobic SSCU increased less steeply than bacterial cell density, possibly as anaerobic metabolism became more important. Across habitats, aerobic SSCU differed significantly in both overall metabolism and diversity of carbon sources. Aerobic metabolism was generally lower for sediment microbes from intertidal on-bottom oyster aquaculture than from eelgrass. Humans indirectly alter microbial activity through biogenic habitats created during aquaculture, but, as has been shown for bivalves more generally, these changes were relatively small, particularly in comparison to sediment changes along estuarine gradients.     

Spatial distribution of fishes and decapods in eelgrass (Zostera marina L.) and sandy habitats of a New Brunswick estuary, eastern Canada

We investigated spatial distribution of fishes and decapods in eelgrass (Zostera marina L.) and non-vegetated sandy habitats in Kouchibouguac Estuary, New Brunswick, Canada. During the ice-free season in 1999 and 2000, mobile fauna were sampled using fyke nets, minnow traps and an epibenthic sled. In general, fishes and decapods were more abundant and diverse (species richness) in eelgrass beds than in nearby sandy habitats, although the trend in abundance of decapods was not statistically significant. Abundance was greater at night than during daylight hours for most species. Atlantic tomcod (Microgadus tomcod), winter flounder (Pleuronectes americanus) and sand shrimp (Crangon septemspinosa) dominated night/crepuscular catches, whereas threespine stickleback (Gasterosteus aculeatus) and cunner (Tautogolabrus adspersus) predominated in diurnal periods. The nursery function of eelgrass habitat was most evident for juvenile white hake (Urophycis tenuis) and small cunners (<3 cm), which were found only in such habitat. This study is one of the few to have investigated the ecological importance of eelgrass habitats for mobile faunal communities in eastern Canada.     

Top–down control by great blue herons Ardea herodias regulates seagrass‐associated epifauna

Top predators can influence the structure and function of plant and animal communities. In coastal marine systems, fish, shark and mammal population declines are major drivers of recent ecosystem‐level change. Cascading effects of predatory wading birds, however, are less understood, even though wading bird populations have declined in many regions. We quantified the effects of predation by the piscivorous great blue heron Ardea herodias fannini on fish, invertebrates and epiphytes living in eelgrass Zostera marina. We found that herons forage on benthic fish in seagrass meadows, and foraging intensity increased from late spring until midsummer. When we experimentally excluded herons, benthic fish abundance increased, and the invertebrate assemblage shifted to more shrimp‐dominated assemblages while grazing gammarid amphipod abundance declined. These shifts were associated with reduced epiphyte abundance when herons were excluded, reflecting a four‐level trophic cascade and mediated by shifts in the grazer assemblage. In summary, we found that a piscivorous wading bird species exerts top down control in a subtidal seagrass ecosystem. Losses and recovery of wading birds could have ecosystem‐level ecological consequences that may need to be considered in the context of concern for overfishing and predator recovery in marine coastal management.     

Structural complexity and fish body size interactively affect habitat optimality

Predator–prey interactions are strongly influenced by habitat structure, particularly in coastal marine habitats such as seagrasses in which structural complexity (SC) may vary over small spatial scales. For seagrass mesopredators such as juvenile fishes, optimality models predict that fitness will be maximized at levels of SC that enhance foraging but minimize predation risk, both of which are functions of body size. We tested the hypothesis that in eelgrass (Zostera marina) habitat, optimal SC for juvenile giant kelpfish (Heterostichus rostratus), an abundant eelgrass mesopredator in southern California, changes through ontogeny. To do this, we quantified eelgrass SC effects on habitat associations, relative predation risk, and foraging efficiency for three size classes of juvenile giant kelpfish. We found that habitat selection differed with fish size: small fish selected dense eelgrass, whereas larger fish selected sparse eelgrass. Small kelpfish experienced the lowest relative predation risk in dense eelgrass but also had higher foraging efficiency in dense eelgrass, suggesting that dense eelgrass is selected by these fish because it minimizes risk and maximizes potential for growth. Surprisingly, larger kelpfish did not experience lower predation risk than small kelpfish. However, larger kelpfish experienced higher foraging efficiency in sparse eelgrass vs. dense eelgrass, suggesting that they select sparse eelgrass to maximize foraging efficiency. Our study highlights that trade-offs between predation risk and foraging can occur within a single habitat type, that studies should consider how habitat value changes through ontogeny, and that seagrass habitat value may be maximal when within-patch variability in SC is high.     

Intertidal community structure differs significantly between substrates dominated by native eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) and adjacent to the introduced oyster <Emphasis Type="Italic">Crassostrea gigas</Emphasis> (Thunberg) in British Columbia,Canada

Eelgrass beds represent important habitats for marine organisms, but are in decline in many coastal areas around the world. On Cortes Island, British Columbia, Canada, oysters coexist regionally with native eelgrass (Zostera marina L.), but eelgrass is typically absent directly seaward of oyster beds (the “below-oyster cobble zone”). We compared assemblage structure of nekton (fish and swimming macroinvertebrates) and epibenthos (macroinvertebrates and macroalgae) between eelgrass bed and below-oyster habitats. We sampled the intertidal zone on Cortes Island at low tide using two methods: quadrats to enumerate epibenthic macroinvertebrates and macroalgae, and beach seines to enumerate fish and swimming macroinvertebrates. Using multivariate analysis of similarity (ANOSIM), we found that the structure of nektonic and epibenthic assemblages associated with below-oyster cobble zones were significantly different from those in eelgrass-beds. Univariate measures showed that nektonic species richness and abundance were significantly higher in eelgrass beds than in below-oyster cobble habitat, whereas epibenthic species richness and abundance were significantly higher in below-oyster habitat. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Trexler     

Importance of Shallow Water Habitats for Demersal Fishes and Decapod Crustaceans in Penobscot Bay, Maine

We characterized demersal fish and decapod crustacean habitats in 14 shallow (< 12m) areas in Penobscot Bay, Maine, by mapping the distribution of subtidal substrata with an acoustic sea bottom profiler. We identified the aquatic vegetation and the fishes and decapods associated with these habitats. Samples of fishes and decapods in each habitat were collected with a small beam trawl. The seabed at most of the stations sampled was composed of a mosaic of substrata. More species of fishes tow^-1 and higher abundances of fishes and decapods were found in stations with vegetation present, particularly eelgrass, Zostera marina. Decapod species richness tow^-1 was similar between vegetated and unvegetated habitats. Shallow habitats in Penobscot Bay were dominated by juvenile fishes and function as nursery areas. The greater species diversity and higher abundances of epibenthic fishes and decapod crustaceans observed in vegetated habitats, particularly beds of Zostera marina, compared with unvegetated areas in Penobscot Bay conform to the hypothesis that increased habitat complexity results in increased species richness and abundance.The first author is also senior author     

Decline of Canada geese (Branta canadensis) and common goldeneye (Bucephala clangula) associated with a collapse of eelgrass (Zostera marina) in a Nova Scotia estuary

Mean numbers of migrant Canada geese (Branta canadensis) in Antigonish Harbour in the southern Gulf of St. Lawrence (Canada) during October to December were similar (approx. 450–500 birds) for the period 1998–2000. Similarly, during this period, geese used two foraging sites. However, in 2001, the average number of birds decreased by half and the primary foraging sites were used only rarely. This coincided with a decline of about 95% in the biomass of roots and rhizomes of eelgrass (Zostera marina) that occurred between October 2000 and 2001. Eelgrass is the principal food of geese in this estuary. In addition, there was a reduction of around 50% in the numbers of common goldeneye (Bucephala clangula), which feed on invertebrates associated with eelgrass. Lower than usual weekly abundances of geese and goldeneye are probably the result of an unusually short residence time in the estuary, rather than a decline in the total number of visiting migrants. We attribute these changes in the distribution and abundance of geese and goldeneyes to the dramatic decline in eelgrass. Electronic Publication     

Congener comparison of native (Zostera marina) and introduced (Z. japonica) eelgrass at multiple scales within a Pacific Northwest estuary

A congener comparison of native (Zostera marina) and introduced (Zostera japonica) eelgrasses was conducted in Willapa Bay, Washington, USA. Along intertidal transects, Z. japonica (0.1–1.5 m above mean lower low water [MLLW]) occurred above Z. marina (<0.6 m MLLW). Both species declined in shoot length at higher elevation, but Z. japonica was always shorter (20 vs. 100 cm) and occurred at higher shoot density (>3,800 vs. <360 m^?2 in Z. marina). Z. japonica exhibited greater seasonal variation in biomass, with increases supported by both sustained asexual reproduction (rhizome branching) and recruitment from seeds (30 vs. <5% in Z. marina). Z. japonica’s successful invasion appears related to small size and high reproductive output, allowing it to spread quickly in a variable and stressful intertidal environment where competitive effects are low. Based on interannual changes in abundance, the native eelgrass has also recently increased in Willapa Bay, and one hypothesis involves “engineering” of suitable habitat at higher tidal elevations by Z. japonica.     

Wintering bird communities in newly-formed wetland in the Yangtze River estuary

We compared wintering bird communities and their habitats among three shoals at Jiuduansha, a newly-formed wetland in the Yangtze River estuary. The highest species richness and diversity were recorded in Shangsha, which is the highest shoal, and the highest abundance and lowest species diversity were recorded in Xiasha, which is the lowest shoal. Shangsha had the largest abundance of perching birds whereas Xiasha was the most abundant in waterbirds. Bird assemblages showed different associations with the different habitat types—perching birds were favored by reed (Phragmites australis) communities, shallow water foragers and dabbling ducks preferred sea-bulrush (Scirpus mariqueter) communities, and moist-soil foragers and gulls showed a preference for bare intertidal zones. All bird assemblages, however, avoided the smooth cordgrass (Spartina alterniflora) communities, which are dominated by an alien invasive plant. The composition of avian communities was related to habitat types at the three shoals. Our results suggest that the newly-formed tidelands can provide suitable habitats for waterbirds and that the lower tidelands can attract more waterfowl than the higher tidelands. Because the shoal with low species diversity could have exclusive bird species, conservation efforts should not concentrate only on the area with high species diversity. The estuarine wetlands should be considered as a whole when conservation strategies are designed. The alien invasive plant should, moreover, be effectively controlled, to provide suitable habitats for birds.     

Biotic resistance to invasion along an estuarine gradient

Biotic resistance is the ability of native communities to repel the establishment of invasive species. Predation by native species may confer biotic resistance to communities, but the environmental context under which this form of biotic resistance occurs is not well understood. We evaluated several factors that influence the distribution of invasive Asian mussels (Musculista senhousia) in Mission Bay, a southern California estuary containing an extensive eelgrass (Zostera marina) habitat. Asian mussels exhibit a distinct spatial pattern of invasion, with extremely high densities towards the back of Mission Bay (up to 4,000 m⁻²) in contrast with near-complete absence at sites towards the front of the bay. We established that recruits arrived at sites where adult mussels were absent and found that dense eelgrass does not appear to preclude Asian mussel growth and survival. Mussel survival and growth were high in predator-exclusion plots throughout the bay, but mussel survival was low in the front of the bay when plots were open to predators. Additional experiments revealed that consumption by spiny lobsters (Panulirus interruptus) and a gastropod (Pteropurpura festiva) likely are the primary factors responsible for resistance to Asian mussel invasion. However, biotic resistance was dependent on location within the estuary (for both species) and also on the availability of a hard substratum (for P. festiva). Our findings indicate that biotic resistance in the form of predation may be conferred by higher order predators, but that the strength of resistance may strongly vary across estuarine gradients and depend on the nature of the locally available habitat.     

Alteration of microbial community composition and changes in decomposition associated with an invasive intertidal macrophyte

Data demonstrating the effects of biological invaders on microbial communities and microbial processes are scarce, especially in marine environments. Research was conducted at Padilla Bay, Washington, to examine the effect that an invasive intertidal eelgrass, Zostera japonica Aschers & Graebn, has on rates of decomposition, microbial community composition, and the possible implications for associated ecosystem processes in this estuarine environment. A series of observational and experimental studies were conducted in beds of Z. japonica, beds of its native congener, Zostera marina, and mixed eelgrass beds. These studies assessed decomposition of invasive and native eelgrass, enumerated bacterial abundance, and examined sole source carbon usage (SSCU) by microbial assemblages. Z. japonica decomposed more rapidly than its native congener throughout the study period although rates of decomposition were variable. Microbial abundance did not differ among different vegetation compositions although differences in SSCU by microbial assemblages were detected among beds of invasive, native, and mixed eelgrass. These results indicate that this abundant invasive species can accelerate rates of decomposition and alter the associative decomposer community, which may lead to higher carbon and nutrient turnover within Padilla Bay.     

Effects of native predators and eelgrass habitat structure on the introduced Asian mussel Musculista senhousia (Benson in Cantor) in southern California

The ability of predators to control the abundance of non-native species has been little explored in marine systems. Native predators may be used to control non-native species or may confer invasion resistance to communities if predation rates on invaders are density-dependent. We studied the response of southern California native predators to the density of Musculista senhousia (Benson in Cantor, 1842), a small, fast growing mussel that has been introduced from Japan to several coastlines worldwide. We performed field experiments to determine if M. senhousia proportional mortality is density-dependent and if eelgrass Zostera marina L. habitat structure influenced mussel density-dependent mortality. We also evaluated the effect of seagrass habitat structure on the aggregative and functional responses of the predatory gastropod Pteropurpura festiva (Hinds, 1844) to Asian mussel density. In the summer of 2002, P. festiva aggregated in plots with high mussel density and was responsible for nearly all predation on M. senhousia. However, M. senhousia proportional mortality was inversely density-dependent at all levels of eelgrass above-ground and below-ground habitat structure. Asian mussel proportional mortality also was inversely density-dependent and was not influenced by eelgrass habitat structure in the spring of 2004 when wading birds were the chief predator of mussels. In contrast to results for mussel proportional mortality, the aggregative and functional responses of P. festiva varied with seagrass habitat structure. P. festiva density increased with Asian mussel density in plots with low simulated habitat structure, but the relationship between P. festiva density and Asian mussel density was parabolic at zero, intermediate and high levels of habitat structure. In field enclosures, P. festiva exhibited a Type I (linear) functional response to Asian mussel density at low levels of eelgrass structure, and a Type II (hyperbolic) functional response to mussel density at high levels of eelgrass structure. Our results and those of others suggest that the degree to which local benthic communities in southern California are resistant to Asian mussel invasion depends on habitat structure, mussel settlement rates, and the density and diversity of predators.     

Estuarine and habitat-related differences in growth rates of young-of-the-year winter flounder (Pseudopleuronectes americanus) and tautog (Tautoga onitis) in three northeastern US estuaries

Instantaneous growth rates of young-of-the-year winter flounder Pseudopleuronectes americanus (Walbaum) (12.0-60.4 mm standard length, SL) and tautog Tautoga onitis (Linnaeus) (21.4-73.8 mm total length, TL) from three estuarine systems in New Jersey (Great Bay-Little Egg Harbor and Navesink River) and Connecticut (Hammonasset River) were used in an attempt to assess the relative quality of selected nominal habitats. A series of short-term field caging experiments were conducted during 1994 and 1995 in: macroalgae (primarily, Ulva lactuca), eelgrass (Zostera marina), unvegetated areas adjacent to macroalgae and eelgrass and tidal creeks in Spartina dominated marsh. Growth rates varied with habitat, estuary and year. Comparisons across nominal habitats within and among estuaries did not show any one habitat with consistently higher growth, and growth was relatively independent of whether a habitat was vegetated or adjacent to vegetation. The growth rates of winter flounder and tautog from the Hammonasset River were not different among habitats in either year of the study. In the Great Bay-Little Egg Harbor, both winter flounder and tautog had higher growth rates in macroalgae with growth in eelgrass varying significantly between years. Conversely, in the Navesink River both species had higher growth rates in eelgrass. Environmental changes associated with temperature and dissolved oxygen appeared to influence growth rates. Winter flounder growth rate and survival was depressed in tidal marsh creeks in the three estuaries and in vegetated macroalgae habitats in the Navesink River where dissolved oxygen levels were often very low (<2 mgl(-1)) for extended periods. In summary, the growth rates of the young-of-the-year of these two species varied temporally and were dependent on the interaction of both the specific estuary and habitat in which the experiments took place. Further, habitat quality, as defined by relative growth rate, was difficult to evaluate because it can be variable and nominal habitat designations are often not sufficient to define the boundaries of a species habitat requirements.     

Patterns of Fish Use and Piscivore Abundance within a Reconnected Saltmarsh Impoundment in the Northern Indian River Lagoon, Florida

Nearly all saltmarshes in east-central, Florida were impounded for mosquito control during the 1960s. The majority of these marshes have since been reconnected to the estuary by culverts, providing an opportunity to effectively measure exchange of aquatic organisms. A multi-gear approach was used monthly to simultaneously estimate fish standing stock (cast net), fish exchange with the estuary (culvert traps), and piscivore abundance (gill nets and bird counts) to document patterns of fish use in a reconnected saltmarsh impoundment. Changes in saltmarsh fish abundance, and exchange of fish with the estuary reflected the seasonal pattern of marsh flooding in the northern Indian River Lagoon system. During a 6-month period of marsh flooding, resident fish had continuous access to the marsh surface. Large piscivorous fish regularly entered the impoundment via creeks and ditches to prey upon small resident fish, and piscivorous birds aggregated following major fish movements to the marsh surface or to deep habitats. As water levels receded in winter, saltmarsh fish concentrated into deep habitats and emigration to the estuary ensued (200% greater biomass left the impoundment than entered). Fish abundance and community structure along the estuary shoreline (although fringed with marsh vegetation) were not analogous to marsh creeks and ditches. Perimeter ditches provided deep-water habitat for large estuarine predators, and shallow creeks served as an alternative habitat for resident fish when the marsh surface was dry. Use of the impoundment as nursery by transients was limited to Mugil cephalus Linnaeus, but large juvenile and adult piscivorous fish used the impoundment for feeding. In conclusion, the saltmarsh impoundment was a feeding site for piscivorous fish and birds, and functioned as a net exporter of forage fish to adjacent estuarine waters.     

Effects of exotic <Emphasis Type="Italic">Spartina alterniflora</Emphasis> on the habitat patch associations of breeding saltmarsh birds at Chongming Dongtan in the Yangtze River estuary,China

Smooth cordgrass (Spartina alterniflora) is one of the most invasive exotic plants of saltmarshes worldwide. To understand the effects of smooth cordgrass invasion on the habitat use and selection by breeding saltmarsh birds, we compared species number and abundance of breeding birds in native reed (Phragmites australis) and smooth cordgrass-invaded habitats (reed-cordgrass mixed habitats and cordgrass monocultures) at Chongming Dongtan in the Yangtze River estuary, China. We further examined the similarity of bird communities in different habitats and habitat selection by dominant bird species. For saltmarsh generalists, species number and abundance did not differ among the habitats. For saltmarsh specialists, species number and abundance did not differ in reed monocultures and reed-cordgrass mixed habitats but were significantly lower in cordgrass monocultures than in reed monocultures and reed-cordgrass mixed habitats. ANOSIM indicated that the difference in bird communities was larger between cordgrass monocultures and the habitats with reed than between the habitats with reed. The saltmarsh specialists preferred reed monocultures, while saltmarsh generalists avoided reed monocultures. Most species indicated no selection (neither preferred nor avoided) on reed-cordgrass mixed habitats, and no species preferred the cordgrass monocultures. The use of cordgrass monocultures by the common saltmarsh birds was negatively related to their body size. This study suggests that the spread of exotic smooth cordgrass has greatly affected the species composition and structure of local bird communities and has been especially disadvantageous to the saltmarsh specialists.     

Habitat specialization of birds in the Czech Republic: comparison of objective measures with expert opinion

Capsule Expert‐based classification of bird species as habitat specialists and as generalists agrees with objective measures of species’ habitat requirements based on large‐scale monitoring data.

Aims To compare habitat specialization of 137 common bird species breeding in the Czech Republic using three different measures and to test their relationships to species’ abundance and habitat associations.

Methods Data on bird abundance and surveyed habitats were collected through a standardized monitoring scheme of common breeding species in the Czech Republic. From these data we calculated a quantitative species specialization index (SSI). Canonical correspondence analysis (CCA) was applied to calculate species’ habitat niche breadth and the level of association of each species to the main habitats. A panel of 11 local bird experts classified each species as habitat generalist or habitat specialist.

Results Species classified as habitat specialists by expert opinion showed higher habitat specialization according to the SSI, as well as according to CCA‐based habitat niche breadth. These species were also more closely associated with one of the main habitat types. These relationships were significant even after controlling for abundance.

Conclusions As expert opinion accords with the level of species’ habitat specialization expressed using two quantitative objective measures, we suggest that these characteristics reflect real interspecific variation in the breadth of habitat requirements in birds. Interspecific differences in habitat specialization are not caused solely by the variability in abundance among species.     

The Role of Eelgrass in Marine Community Interactions and Ecosystem Services: Results from Ecosystem-Scale Food Web Models

Eelgrass beds provide valuable refuge, foraging, and spawning habitat for many marine species, including valued species such as Pacific salmon (Oncorhynchus spp.), Pacific herring (Clupea pallasi), and Dungeness crab (Metacarcinus magister). We used dynamic simulations in a food web model of central Puget Sound, Washington, USA developed in the Ecopath with Ecosim software, to examine how the marine community may respond to changes in coverage of native eelgrass (Zostera marina), and how these modeled responses can be assessed using an ecosystem services framework, expressing these services with economic currencies in some cases and biological proxies in others. Increased eelgrass coverage was most associated with increases in commercial and recreational fishing with some small decreases in one non-market activity, bird watching. When we considered ecosystem service categories that are aggregations of individual groups of species, we saw little evidence of strong tradeoffs among marine resources; that is, increasing eelgrass coverage was essentially either positive or neutral for all services we examined, although we did not examine terrestrial activities (for example, land use) that affect eelgrass coverage. Within particular service categories, however, we found cases where the responses to changes in eelgrass of individual groups of species that provide the same type of ecosystem service differed both in the magnitude and in the direction of change. This emphasizes the care that should be taken in combining multiple examples of a particular type of ecosystem service into an aggregate measure of that service.     

Twentyfive years of changes in the distribution and biomass of eelgrass,Zostera marina,in Grevelingen lagoon,The Netherlands

The wax and wane of the eelgrass (Zostera marina L.) population in Grevelingen lagoon (East Atlantic; The Netherlands) has been documented for over 25 years, together with quantitative and semi-quantitative data on environmental variables. The population expanded after the closure of the Grevelingen estuary in 1971, but declined from 4600 ha surface area in 1978 to less than 100 ha in 1993. There is little causal evidence which factors are responsible for the observed dynamics of the population. The incomplete picture emerging from the data is that of an extremely impoverished eelgrass population, living under constant oligo-mesotrophic marine conditions. Both the sexual and the vegetative modes of reproduction are severely stressed by environmental variables, most likely a combination of low temperatures, high salinity, low dissolved silicate and low ammonium concentrations. Survival of the population asks for the restoration of moderate estuarine conditions.Contribution No. 2180 of the Netherlands Institute of Ecology, Nieuwersluis, The Netherlands.     

Detecting the impacts of notorious invaders: experiments versus observations in the invasion of eelgrass meadows by the green seaweed <Emphasis Type="Italic">Codium fragile</Emphasis>

Biological invasions can vary in the extent of their effects on indigenous communities but predicting impacts for particular systems remains difficult. In coastal marine ecosystems, the green seaweed Codium fragile ssp. fragile is a notorious invader with its reputation based on studies conducted largely on rocky shores. The green seaweed has recently invaded soft-bottom eelgrass communities by attaching epiphytically to eelgrass (Zostera marina) rhizomes, thereby creating the potential for disruption of these coastal habitats through competition or disturbance. We investigated the effect of this invader on various aspects of eelgrass performance (shoot density and length, shoot growth, above- and below-ground biomass, carbohydrate storage) using both small-scale manipulative and large-scale observational experiments. Manipulative experiments that varied Codium abundance demonstrated clear negative effects over a 4-month period on shoot density and carbohydrate reserves, but only for high, but realistic, Codium biomass levels. Light levels were much lower under canopies for high and medium density Codium treatments relative to low and control Codium cover treatments, suggesting that shading may influence eelgrass growing under the algal cover. In contrast, these effects were either not detectable or very weak when examined correlatively with field surveys conducted at larger spatial scales, even for sites that had been invaded for over 4 years. It is premature to extend generalizations of Codium’s impact derived from studies in other systems to eelgrass communities; further efforts are required to assess the long-term threats that the alga poses to this ecosystem. This study demonstrates the need to investigate impacts of invasions over multiple scales, especially those that incorporate the temporal and spatial heterogeneity of the invader’s abundance.     

Formal analysis and evaluation of allometric methods for estimating above‐ground biomass of eelgrass

Eelgrass (Zostera marina) populations supply substantial amounts of organic materials to food webs in shallow coastal environments, provide habitat for many fishes and their larvae and abate erosion. The characterisation of eelgrass biomass dynamics is an important input for the assessment of the function and values for this important seagrass species. We here present original allometric methods for the non‐destructive estimation of above‐ground biomass of eelgrass. These assessments are based on measurements of lengths and areas of leaves and sheaths and mathematical models that can be identified by means of standard regression procedures. The models were validated by using data obtained from Z. marina meadows in the Punta Banda estuary B.C., Mexico, and in Jindong Bay, Korea. Using available data and concordance correlation index criteria we show that the values projected thorough the presented allometric paradigm reproduces observed values in a consistent way. The annual average value for observed above‐ground biomass was 1.46 ± 0.15 g shoot^?1, while the corresponding calculated value was 1.40 ± 0.13 g shoot^?1. We suggest that our method can be applied to other studies in which the architecture and growth form of leaves and sheaths are similar to those of eelgrass. This would provide reliable and simplified estimations of biomass while eliminating tedious laboratory processing and avoiding destructive sampling.