Salt marsh shoreline geomorphology influences the success of restored oyster reefs and use by associated fauna

Restoration is increasingly implemented as a strategy to mitigate global declines in biogenic habitats, such as salt marshes and oyster reefs. Restoration efforts could be improved if we knew how site characteristics at landscape scales affect the ecological success of these foundation species. In this study, we determined how salt marsh shoreline geomorphologies (e.g. with variable hydrodynamic energy, fetch, erosion rates, and slopes) affect the success of restored intertidal oyster reefs, as well as how fauna utilize restored reefs and forage along marsh habitats. We constructed oyster reefs along three marsh shoreline geomorphologies in May 2012: 1) “creek” (small‐fetch, gradual‐sloped shoreline), “ramp” (large‐fetch, gradual‐sloped shoreline), and “scarp” (large‐fetch, steep‐sloped shoreline). Following recruitment, oyster spat density was greatest on ramp reefs; however, 2 years later, the highest adult oyster densities were found on creek reefs. Total nekton and blue crab catch rates in trawl nets were highest in the creek, while piscivore catch rates in gill nets were highest along the scarp shoreline. We found no difference in predation on snails in the salt marsh behind constructed reef and nonconstructed reference sites, but there were more snails consumed in the creek shoreline, which corresponded with the distribution of their major predator—blue crabs. We conclude that oyster reef construction was most successful for oysters in small‐fetch, gradual‐sloped, creek environments. However, nekton abundance did not always follow the same trends as oyster density, which could suggest constructed reefs may offer similar habitat‐related functions (prey availability and refuge) already present along existing salt marsh borders.     

Comparisons of macrofaunal assemblages on restored and non-restored oyster reefs in mesohaline regions of Chesapeake Bay in Maryland

Maryland's recently created oyster restored reefs provide us with a unique opportunity to observe the abundance and species composition of macrofauna assemblages on unexploited reefs with high concentrations of mature oysters and undisturbed reef architecture. They might thus be used to better understand the magnitude of losses to reef dwelling macrofauna communities, and the associated loss of ecological functions resulting from reef destruction. We sampled reef macrofaunal assemblages on restored plots at four restored oyster reefs and adjacent non-restored plots located outside restored boundaries. We then compared the effects of study site location, and habitat quality (restored versus non-restored) on macrofaunal density using thirteen response variables. Density of macrofauna was an order of magnitude higher on restored reefs, epifaunal density was more than twice as high on restored reefs and sessile macrofaunal density was two orders of magnitude higher on restored reefs. Three out of the five dominant taxonomic groups were much more abundant on restored plots. Mean amphipod density was 20 times higher on restored plots and densities of xanthid crabs and demersal fish were both four times greater on restored plots. Two out of four functional feeding groups: suspension feeders and carnivore/omnivores, were more abundant on restored plots. Since reef macrofauna include many important fish prey species, oyster reef restoration may have the potential to augment fish production by increasing fish prey densities and fish foraging efficiency.     

The Eocene fishes of Monte Bolca: the earliest coral reef fish assemblage

The fish assemblage from the Eocene deposits of Monte Bolca, Northern Italy, are compared with those of Recent coral reefs. A family-level taxonomic definition of a Recent coral reef fish assemblage is formulated to permit direct comparisons. On this basis, the Monte Bolca fishes represent the earliest clearly defined coral reef fish assemblage. Quantitative analyses of the relative abundance of fish families revealed significant differences between the two assemblages. The Bolca assemblage has Mesozoic links (Pycnodontiformes) and non-perciform taxa are relatively abundant, particularly the Beryciformes (Holocentridae). However, Bolca represents the earliest record of a perciform-dominated benthic fish assemblage (68.4% of all non-clupeid taxa). Within the Perciformes, the abundance of the major reef fish lineages (higher squamipinnes and Labroidei) differs markedly between the two assemblages. The numerical dominance of labroid fishes on coral reefs appears to have been a relatively recent occurrence.     

Distribution and environmental control of coral assemblages in northern Safaga Bay (Red Sea, Egypt)

Summary Coral assemblages in northern Safaga Bay, Red Sea, Egypt, are qualitatively described. Nine distinct assemblages were found, which correspond to quantitatively defined community types previously described from the area off Hurghada, northern Red Sea. Their distribution within northern Safaga Bay was mapped. Strong gradient and/or steep relief assemblages were:Acropora assemblage on windward (exposed) reefs,Porites assemblage on leeward (sheltered) reefs,Millepora assemblage on current exposed reefs,Stylophora assemblage on reef flats. Low gradient and/or low relief assemblages were:Acropora dominated coral patches in areas of good circulation to a depth of 15 m,Stylophora/Acropora coral patch assemblages in shallow sheltered environments, faviid carpet in low relief areas between 10 and 25 m which with increasing turbidity turns into a depauperate faviid carpet,Porites carpet in low relief areas between 5 and 15 m with clearest water,Sarcophyton carpet in low relief areas with high suspension load, platy scleractinian assemblage in deeper water (>25 m) with low light intensity. The distribution of coral assemblages depends basically on 1) topography 2) hydrodynamics 3) light and 4) suspension load.     

河北唐山曹妃甸-乐亭海域自然牡蛎礁分布及生态意义

牡蛎礁是生态系统服务价值最高的海洋生境之一,目前我国自然牡蛎礁分布和生态现状的基础信息仍然较缺乏。于2019年3月对河北唐山曹妃甸-乐亭海域自然牡蛎礁的空间分布、生态环境、牡蛎生物学和礁体动物群落开展了调查,并评估了该牡蛎礁的生态系统服务价值。该海域自然牡蛎礁分布于溯河(SR)、溯河口海域(SRE)和捞鱼尖海域(LYJ),总面积约15 km~2,是目前我国面积最大的自然牡蛎礁。基于96个牡蛎样品的16S rDNA检测,共识别出92个长牡蛎Crassostrea gigas、3个侏儒牡蛎Nanostrea fluctigera和1个巨蛎属未知种Crassostrea sp.。自然牡蛎礁中牡蛎平均密度介于104—3912个/m~2之间,不同礁区间牡蛎平均密度的大小排序为：SRE>SR>LYJ (P<0.05),平均生物量的大小排序为：SR>SRE>LYJ (P<0.05),平均壳高的大小排序为：SR>SRE=LYJ (P<0.05)。在该牡蛎礁内记录到49种礁体动物,其中软体动物16种、节肢动物16种、环节动物8种、棘皮动物5种、腔肠动物2种...     

Herbivorous fish assemblages and herbivory pressure on Laamu Atoll, Republic of Maldives

While herbivory is recognized as a fundamental process structuring coral reef communities, herbivore assemblages and processes are poorly described for reefs in the Indian Ocean region. We quantified herbivorous fish assemblage structure (abundance and diversity) in Laamu Atoll, Republic of Maldives, in four reef habitat types: faro reef flats, faro reef slopes, inner and outer atoll reef slopes (20 sites in total). Herbivorous fish assemblages, representing a total of 30 species, grouped strongly by habitat type, with the highest absolute abundance observed on faro reef flats and lowest abundance on inside atoll rim reef slopes. Removal of Thalassia seagrass blades by ambient herbivore assemblages was used in a bioassay to assess relative herbivory pressure among four habitat types (eight sites). Also, at one site a choice herbivory assay was performed to assess herbivore preference among four benthic plants across three depth zones. Relative herbivory, as indicated by Thalassia assays, was highest on inside atoll rim reef slopes and lowest on outside atoll rim reef slopes. Thalassia consumption did not correspond to overall herbivorous fish abundance, but corresponded more closely with parrotfish abundance. In the choice assays, herbivores showed strong preferences among plant types and consumption of most plant types was higher at mid-depth than in the shallow reef flat or deep reef knoll zones.     

Evaluating Intertidal Oyster Reef Development in South Carolina Using Associated Faunal Indicators

Eastern oyster (Crassostrea virginica) habitat is increasingly being restored for the ecosystem services it provides rather than solely as a fishery resource. Community‐based projects with the goal of ecological restoration have successfully constructed oyster reefs; however, the habitat benefits of these restoration efforts are usually not assessed or reported. In this study, we examined oyster habitat development at five community‐based oyster restoration sites in South Carolina using oyster population parameters, resident fauna densities, and sedimentation (percent sediment coverage) as assessment metrics. All sites included multiple‐aged reefs (1–3 years old) at the time of the fall 2004 sampling. Resident crabs and mussels were abundant at all five sites and crab assemblages were related to the size structure of the oyster microhabitat. Scorched mussel (Brachidontes exustus) abundances were most frequently correlated with oyster and other resident species abundances. Associations among oysters and resident crabs and mussels were not evident when analyses were conducted with higher level taxonomic groupings (e.g., total number of crabs, mussels, or oysters), indicating that species‐level identifications improve our understanding of interactions among reef inhabitants and oyster populations. Community‐based restoration sites in South Carolina provide habitat for mussels and resident crabs, in some cases in the absence of dense populations of relatively large oysters. Monitoring programs that neglect species‐level identifications and counts of mussels and crabs may underestimate the successful habitat provision that can arise independent of large, dense oyster assemblages.     

Predator size interacts with habitat structure to determine the allometric scaling of the functional response

While both predator body size and prey refuge provided by habitat structure have been established as major factors influencing the functional response (per capita consumption rate as a function of prey density), potential interactions between these factors have rarely been explored. Using a crab predator (Panopeus herbstii) – mussel prey (Brachidontes exustus) system, we examined the allometric scaling of the functional response in oyster (Crassostrea virginica) reef habitat, where crevices within oyster clusters provide mussels refuge from predation. A field survey of mussel distribution showed that mussels attach closer to the cluster periphery at high mussel density, indicating the potential for saturation of the refuge. In functional response experiments, the consumption rate of large crabs was depressed at low prey density relative to small crabs, while at high prey density the reverse was true. Specifically, the attack rate coefficient and handling time both decreased non‐linearly with crab size. An additional manipulation revealed that at low prey densities, the ability of large crabs to maneuver their claws and bodies to extract mussels from crevices was inhibited relative to small crabs by the structured habitat, reducing their attack rate. At high prey densities, crevices were saturated, forcing mussels to the edge of clusters where crabs were only limited by handling time. Our study illuminates a potentially general mechanism where the quality of the prey refuge provided by habitat structure is dependent on the relative size of the predator. Thus anthropogenic influences that alter the natural crab size distribution or degrade reef habitat structure could threaten the long‐term stability of the crab –mussel interaction in reefs.     

Structure of Caribbean coral reef communities across a large gradient of fish biomass

The collapse of Caribbean coral reefs has been attributed in part to historic overfishing, but whether fish assemblages can recover and how such recovery might affect the benthic reef community has not been tested across appropriate scales. We surveyed the biomass of reef communities across a range in fish abundance from 14 to 593 g m⁻², a gradient exceeding that of any previously reported for coral reefs. Increased fish biomass was correlated with an increased proportion of apex predators, which were abundant only inside large marine reserves. Increased herbivorous fish biomass was correlated with a decrease in fleshy algal biomass but corals have not yet recovered.     

Facies and palaeoecology of Upper Jurassic (Middle Oxfordian) coral reefs in England

Summary This study documents the facies and fauna of Late Jurassic (Middle Oxfordian) coral reefs in England. Sedimentological and palaeoecological analysis of these reefs distinguishes three generic reef types: (1) small reef patches and thickets associated with siliciclastic deposits; (2) small reef patches and thickets associated with siliciclastic-free bioclastic grainstones and packstones; and (3) biostromal units associated with deep water facies. The depositional environments of these reef types are discussed. Two coral assemblages are identified: (1) the microsolenid assemblage; and (2) theThamnasteria, Isastraea, Fungiastraea andThecosmilia assemblage (Thamnasteria assemblage). TheThamnasteria assemblage developed in all shallow water environments in the study area, regardless of local environmental conditions. The fauna is very eurytopic,r-selected and can tolerate significant environmental fluctuations on short temporal scales (sub-seasonal). The main control on the development of the microsolenid assemblage was low light intensity, low background sedimentation rates and low hydrodynamic energy levels.     

Energy flow to two abundant consumers in a subtropical oyster reef food web

Oyster reefs are among the most threatened coastal habitat types, but still provide critical habitat and food resources for many estuarine species. The structure of oyster reef food webs is an important framework from which to examine the role of these reefs in supporting high densities of associated fishes. We identified major trophic pathways to two abundant consumers, gray snapper (Lutjanus griseus) and crested goby (Lophogobius cyprinoides), from a subtropical oyster reef using stomach content and stable isotope analysis. The diet of gray snapper was dominated by crabs, with shrimp and fishes also important. Juvenile gray snapper fed almost entirely on oyster reef-associated prey items, while subadults fed on both oyster reef- and mangrove-associated prey. Based on trophic guilds of the gray snapper prey, as well as relative δ¹³C values, microphytobenthos is the most likely basal resource pool supporting gray snapper production on oyster reefs. Crested goby had omnivorous diets dominated by bivalves, small crabs, detritus, and algae, and thus were able to take advantage of prey relying on production from sestonic, as well as microphytobenthos, source pools. In this way, crested goby represent a critical link of sestonic production to higher trophic levels. These results highlight major trophic pathways supporting secondary production in oyster reef habitat, thereby elucidating the feeding relationships that render oyster reef critical habitat for many ecologically and economically important fish species.     

Regional and local drivers of macroinvertebrate assemblages in boreal springs

Aim To identify the most important environmental drivers of benthic macroinvertebrate assemblages in boreal springs at different spatial scales, and to assess how well benthic assemblages correspond to terrestrially derived ecoregions. Location Finland. Methods Benthic invertebrates were sampled from 153 springs across four boreal ecoregions of Finland, and these data were used to analyse patterns in assemblage variation in relation to environmental factors. Species data were classified using hierarchical divisive clustering (twinspan ) and ordinated using non‐metric multidimensional scaling. The prediction success of the species and environmental data into a priori (ecoregions) and a posteriori (twinspan ) groups was compared using discriminant function analysis. Indicator species analysis was used to identify indicator taxa for both a priori and a posteriori assemblage types. Results The main patterns in assemblage clusters were related to large‐scale geographical variation in temperature. A secondary gradient in species data reflected variation in local habitat structure, particularly abundance of minerogenic spring brooks. Water chemistry variables were only weakly related to assemblage variation. Several indicator species representing southern faunistic elements in boreal springs were identified. Discriminant function analysis showed poorer success in classifying sites into ecoregions based on environmental than on species data. Similarly, when classifying springs into the twinspan groups, classification based on species data vastly outperformed that based on environmental data. Main conclusions A latitudinal zonation pattern of spring assemblages driven by regional thermal conditions is documented, closely paralleling corresponding latitudinal patterns in both terrestrial and freshwater assemblages in Fennoscandia. The importance of local‐scale environmental variables increased with decreasing spatial extent. Ecoregions provide an initial stratification scheme for the bioassessment of benthic macroinvertebrates of North European springs. Our results imply that climate warming, landscape disturbance and degradation of spring habitat pose serious threats to spring biodiversity in northern Europe, especially to its already threatened southern faunistic elements.     

Land‐use legacy and the differential response of stream macroinvertebrates to multiple stressors studied using in situ experimental mesocosms

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Fish assemblages on sunken vessels and natural reefs in southeast Florida,USA

Derelict ships are commonly deployed as artificial reefs in the United States, mainly for recreational fishers and divers. Despite their popularity, few studies have rigorously examined fish assemblages on these structures and compared them to natural reefs. Six vessel-reefs off the coast of southeast Florida were censused quarterly (two ships per month) to characterize their associated fish assemblages. SCUBA divers used a non-destructive point-count method to visually assess the fish assemblages over 13- and 12-month intervals (March 2000 to March 2001 and March 2002 to February 2003). During the same intervals, fish assemblages at neighboring natural reefs were also censused. A total of 114,252 fishes of 177 species was counted on natural and vessel-reefs combined. Mean fish abundance and biomass were significantly greater on vessel-reefs in comparison to surrounding natural reef areas. Haemulidae was the most abundant family on vessel-reefs, where it represented 46% of total fish abundance. The most abundant family on natural reefs was Labridae, where it accounted for 24% of total fish abundance. Mean species richness was significantly greater on vessel-reefs than neighboring natural reefs and also differed among vessel-reefs. Both mean fish abundance and mean species richness were not significantly different between natural reefs neighboring vessel-reefs and natural reefs with no artificial structures nearby. This suggests the vessel-reefs are not, in general, attracting fish away from neighboring natural reefs in our area. Additionally, economically important fish species seem to prefer vessel-reefs, as there was a greater abundance of these species on vessel-reefs than surrounding natural reef areas. Fish assemblage structure on natural versus artificial reefs exhibited a low similarity (25.8%). Although no one species was responsible for more than 6% of the total dissimilarity, fish assemblage trophic structure differed strikingly between the two reef types. Planktivores dominated on vessel-reefs, accounting for 54% of the total abundance. Conversely, planktivores only made up 27% of total abundance on natural reefs. The results of this study indicate vessel-reef fish assemblages are unique and that these fishes may be utilizing food resources and habitat characteristics not accessible from or found at natural reefs in our area. Production may also be occurring at vessel-reefs as the attraction of fish species from nearby natural reefs seems to be minimal. Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users     

Do restored oyster reefs benefit seagrasses? An experimental study in the Northern Gulf of Mexico

Oyster reefs and seagrass beds are being lost worldwide at alarming rates. These habitats provide many services to humankind and, thus, much effort has been dedicated to their restoration. Here, we examine the efficacy of created oyster reefs at enhancing seagrass beds through the amelioration of hydrographic conditions and water quality. We carried out a field experiment in the Northern Gulf of Mexico where we compared areas shoreward of created reefs with adjacent reef‐free areas over several years using a before‐after control‐impact (BACI) design. The reefs were built with oyster shell, measured 65 m, and were placed at circa 100 m from the shoreline to ensure subtidal conditions and enhance oyster recruitment. The BACI results showed few and disparate effects of the reefs, even when distance from the reef was factored in. However, we found a temporal increase in seagrass cover throughout all the experimental area (i.e. including both reef and control plots) following reef deployment. Interestingly, further analysis with satellite imagery showed the experimental area had higher seagrass cover 5 years after reef deployment than it did before reef deployment, but such increase was not observed for nearby areas. In concert, the results suggest “shadow” effects for the reefs examined, where positive effects on seagrass beds extend beyond the area directly shoreward from the reef. Oyster reef restoration may have positive impacts on shallow seagrass beds in turbid, high‐energy systems; however, more work on the extent and mechanisms for this interaction is needed.     

Faunal utilization of constructed intertidal oyster (Crassostrea rivularis) reef in the Yangtze River estuary,China

An intertidal oyster reef (～260 ha) was created by planting hatchery-reared seed oysters (Crassostrea rivularis) on an artificial concrete modular reef in the Deepwater Navigation Channel Regulation Project of the Yangtze River estuary. We examined the development of reef communities (oyster, barnacle and motile epibenthic macrofauna), characterized nekton use and assessed the habitat value of the constructed reef. The C. rivularis oyster population showed a rapid exponential increase with time and reached maximum density (3410 ± 241 ind./m²) and biomass (3175 ± 532 g/m²) after one year of restoration. The barnacle Balanus albicostatus was the most abundant sessile macrofauna and had a significantly greater density in the high intertidal zone than in the low intertidal zone (P < 0.05). The reef also supported diverse motile epibenthic macrofauna (11 mollusks, 11 crustaceans, 4 annelids and 2 fishes), and the reef-associated communities were numerically dominated by Neanthes japonica, Perinereis aibuhitensis, Nerita yoldi and Littorinopsis intermedia. A total of 50 nekton species (31 fishes, 9 shrimps and 10 crabs) utilized the constructed intertidal oyster reef, and grass shrimp Palaemon spp. dominated the nekton communities in term of abundance. Since the constructed intertidal oyster reef supports a variety of reef communities and abundant nektons, it should be recognized as an important and protective fish habitat in the Yangtze River estuary.     

Effects of Reef Proximity on the Structure of Fish Assemblages of Unconsolidated Substrata

Fish assemblages of unconsolidated sedimentary habitats on continental shelves are poorly described when compared to those of hard substrata. This lack of data restricts the objective management of these extensive benthic habitats. In the context of protecting representative areas of all community types, one important question is the nature of the transition from reefal to sedimentary fish assemblages. We addressed this question using Baited Remote Underwater Videos (BRUVs) to assess fish assemblages of sedimentary habitats at six distances from rocky reefs (0, 25, 50, 100, 200, and 400 m) at four sites in subtropical eastern Australia. Distance from reef was important in determining fish assemblage structure, and there was no overlap between reef sites and sedimentary sites 400 m from reef. While there was a gradient in assemblage structure at intermediate distances, this was not consistent across sites. All sites, however, supported a mixed ‘halo’ assemblage comprising both reef and sediment species at sampling stations close to reef. BRUVs used in conjunction with high-resolution bathymetric and backscatter spatial data can resolve differences in assemblage structure at small spatial scales (10s to 100s of metres), and has further application in unconsolidated habitats. Unless a ‘reef halo’ assemblage is being examined, a minimum of 200 m but preferably 400 m distance from any hard substrate is recommended when designing broader-scale assessments of fish assemblages of sedimentary habitats.     

An invasive crab alters interaction webs in a marine community

Over the last decade, the non-native, filter-feeding crab Petrolisthes armatus invaded oyster reefs of the South Atlantic Bight at densities of thousands m⁻². Mesocosm and field experiments demonstrated that P. armatus at ∼10–75% of mean summer densities: (1) suppressed growth of small oysters, biomass of benthic microalgae, and recruitment of native mud crabs, (2) enhanced oyster, mussel, and total bivalve recruitment, macroalgal cover, and survivorship of predatory oyster drills, but (3) did not affect native taxonomic richness. Laboratory feeding assays, field tethering experiments, and population changes in field and mesocosm experiments suggest that P. armatus is a preferred prey for native mud crabs and other consumers, thus relieving predation on native species and enhancing recruitment or survival of bivalves and oyster drills. In contrast, the invasive crab can consume crustacean larvae and via this feeding may suppress recruitment of native mud crabs. Our findings should be conservative given the low densities of P. armatus seeded into experimental plots and our inability to run longer-term experiments due to controls rapidly being colonized by non-native crabs recruiting from the plankton. Invasive crabs commonly impact native communities via predation, but community impacts of this invasive crab may be as much due to its role as a preferred prey of native consumers as to its predation on native prey. Given that oysters are foundation species for shallow reefs in the South Atlantic Bight, the long-term effects of this invasion could be considerable.     

Spatial variation of reef fishes and the relative influence of biotic and abiotic habitat traits

Patterns of distribution of reef fishes were examined across three spatial scales and related to habitat traits along 25 km of the northern Portuguese coast. Response variables included the multivariate assemblage structure, the total number of taxa and individuals, and the abundance of single groups categorized according to their preference for the benthic, proximo-benthic or pelagic environment, feeding and reproductive behaviour. Habitat traits included topographic elements (small and large ‘drops’ like cracks and crevices) and the extent of dominant morpho-functional types of macroalgae (kelp, large foliose, small erect, turf-forming filamentous, and encrusting). All fish responses were characterized by the largest variance at the smallest scale (among transects tens m apart), followed by that among reefs (hundreds m to 1 km apart) and almost null variance among sites (some km apart). Small and large ‘drops’ of the substratum explained, respectively, considerable variation of assemblage structure and the total abundance of individuals, while the extent of bare rock influenced the richness of taxa and that of benthic fishes, fishes feeding on sessile invertebrates and fishes laying benthic eggs or having nesting behaviour. Combinations of abiotic and biotic structural attributes of reefs influenced proximo-benthic fishes, the predators of mobile animals and fishes releasing pelagic eggs. The here reported associations between patterns of distribution of reef fishes and habitat traits have implications for the design of future protection schemes suitable to guarantee the conservation of reef fish communities and of the processes responsible for their variation. Within the SLOSS (single-large vs. several-small) debate in the design of marine reserves, for example, effective protection to the studied reef fishes would be provided by a set of small reserves, rather than a single large which might be appropriate for fishes having wider home ranges.