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.     

Cuing oyster recruitment with shell and rock: implications for timing reef restoration

Reducing uncertainty surrounding the biological responses of degraded habitat is key to providing confidence and efficiency in its restoration. Many coastal habitats are so extensively degraded that organismal responses to restoration interventions are entirely unknown. Among the most degraded coastal ecosystems are oyster reefs, whose restoration typically occurs where oysters are degraded to the point of functional extinction. This loss creates uncertainty on the fundamental processes for reef recovery; the timing of oyster recruitment and their preferred substratum for settlement. Such knowledge can inform restoration strategies to accelerate habitat recovery. Near the site of Australia's largest restoration of native oyster reef, we quantified temporal variability in recruitment of the native flat oyster (Ostrea angasi) and assessed their preference between the settlement substrata deployed for the reef restoration. Combining half a decade of environmental data with oyster recruitment data, we provide a model that identified distinct peaks in oyster recruitment which correlate with food availability and seawater temperature. In addition, oysters preferentially settled on oyster shell relative to other materials used in local restoration. In combination, these results suggest that there are opportunities to augment recruitment through addition of shell substratum synced with recruitment peaks. Our recruitment model likely represents a minor investment with large returns, providing opportunities to capture peak recruitment and greater confidence in utilizing natural recruitment as a restoration resource.     

Disturbance and reef topography maintain high local diversity in Ecklonia radiata kelp forests

Disturbance of competitive‐dominant plant and algae canopies often lead to increased diversity of the assemblage. Kelp forests, particularly those of temperate Western Australia, are habitats with high alpha diversity. This study investigated the roles of broad‐scale canopy loss and local scale reef topography on structuring the kelp‐dominated macroalgal forests in Western Australia. Eighteen 314 m² circular areas were cleared of their Ecklonia radiata canopy and eighteen controls were established across three locations. The patterns of macroalgal recolonisation in replicate clearances were observed over a 34 month period. Macroalgal species richness initially increased after canopy removal with a turf of filamentous and foliose macroalgae dominating cleared areas for up to seven months. A dense Sargassum canopy dominated cleared areas from 11 to 22 months. By 34 months, partial recovery of the kelp canopy into cleared areas had occurred. Some cleared areas did not follow this trajectory but remained dominated by turfing, foliose and filamentous algae. As kelp canopies developed, the initial high species diversity declined but still remained elevated relative to undisturbed controls, even after 34 months. More complex reef topography was associated with greater variability in the algal assemblage between replicate quadrats suggesting colonising algae had a greater choice of microhabitats available to them on topographically complex reefs. Shading by canopies of either Sargassum spp. and E. radiata are proposed to highly influence the abundance of algae through competitive exclusion that is relaxed by disturbance of the canopy. Disturbance of the canopy in E. radiata kelp forests created a mosaic of different patch types (turf, Sargassum‐dominated, kelp‐dominated). These patch types were both transient and stable over the 34 months of this study, and are a potential contemporary process that maintains high species diversity in temperate kelp‐dominated reefs.     

Accounting for Multiple Foundation Species in Oyster Reef Restoration Benefits

Many coastal habitat restoration projects are focused on restoring the population of a single foundation species to recover an entire ecological community. Estimates of the ecosystem services provided by the restoration project are used to justify, prioritize, and evaluate such projects. However, estimates of ecosystem services provided by a single species may vastly under‐represent true provisioning, as we demonstrate here with an example of oyster reefs, often restored to improve estuarine water quality. In the brackish Chesapeake Bay, the hooked mussel Ischadium recurvum can have greater abundance and biomass than the focal restoration species, the eastern oyster Crassostrea virginica. We measured the temperature‐dependent phytoplankton clearance rates of both bivalves and their filtration efficiency on three size classes of phytoplankton to parameterize an annual model of oyster reef filtration, with and without hooked mussels, for monitored oyster reefs and restoration scenarios in the eastern Chesapeake Bay. The inclusion of filtration by hooked mussels increased the filtration capacity of the habitat greater than 2‐fold. Hooked mussels were also twice as effective as oysters at filtering picoplankton (1.5–3 µm), indicating that they fill a distinct ecological niche by controlling phytoplankton in this size class, which makes up a significant proportion of the phytoplankton load in summer. When mussel and oyster filtration are accounted for in this, albeit simplistic, model, restoration of oyster reefs in a tributary scale restoration is predicted to control 100% of phytoplankton during the summer months.     

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...     

Effects of time and harvest on genetic diversity of natural and restored oyster reefs

Oysters serve as keystone species and ecosystem engineers in estuaries due to their fundamental role of providing services to the surrounding environment and to humans. Globally, however, oysters have precipitously declined in numbers over the last century. To remedy this drastic decrease, many coastal areas have initiated oyster restoration projects. In the Indian River Lagoon (IRL) of Florida, where oyster loss was primarily the result of recreational boat wake dislodgment, researchers have put in place a unique method to supplement natural recruitment of oysters. This method consists of deploying stabilized shell attached to mesh mats. Larval oysters thus have substrate on which to settle and three‐dimensional reef habitats have been reestablished in historical footprints. This restoration project has proven to be successful, shown by 9 years of data collection on growth, recruitment, and survivability. In this study, we sought to determine the length of time required for newly restored oyster reefs to reach equivalent levels of genetic diversity as undisturbed, natural (reference) oyster reefs. Additionally, we determined if recreational harvesting impacted the genetic diversity of these reference reefs. Using nine microsatellite loci, we found that restored oyster reefs accumulated as much genetic diversity as natural reefs as quickly as 1 month after stabilized shells were deployed. We likewise found that harvesting did not impact genetic diversity in oyster reefs in the IRL. These results are encouraging, and are a valuable addition to understanding the importance of oyster reef restoration on the ecosystem.     

Oyster reef restoration: substrate suitability may depend on specific restoration goals

A limited supply of oyster shell for restoration practices has prompted investigations of alternative substrates used in construction of artificial oyster reefs. The success of oyster reef restoration projects is increasingly focused not only on oyster densities, but also on habitat provisioning for associated fauna. A subtidal oyster reef complex (0.24 km²) was restored in the Mission‐Aransas Estuary, Texas, U.S.A., in July 2013 using replicated mounds of concrete, limestone, river rock, and oyster shell substrates. Oyster and reef‐associated fauna characteristics were quantified quarterly for 15 months, using sampling trays that were deployed 3 months after construction. The highest densities of oyster spat occurred 9 months after tray deployment (July 2014, 1,264/m²), whereas juvenile oyster densities increased throughout the study period to 283/m². Concrete (1,022/m²) and limestone (939/m²) supported the highest number of oysters over all dates. Oyster shell (1,533/m²) and concrete (1,047/m²) substrates supported the highest densities of associated motile fauna. Faunal diversity (Hill's N1) did not vary by substrate material, but did show seasonal variation. A simple benefit–cost ratio was used to indicate the localized monetary value for each of the substrates. Oyster shell and concrete substrates returned the highest benefit–cost ratio for motile fauna, while concrete yielded the highest benefit–cost ratio for oyster abundance. Incorporating benefit–cost ratios in restoration planning will allow practitioners to better integrate substrate‐specific ecological values with economic considerations and project goals to maximize return on restoration investments.     

Predator-prey dynamics between recently established stone crabs (Menippe spp.) and oyster prey (Crassostrea virginica)

Range expansion and population establishment of individual species can have significant impacts on previously established food webs and predator-prey dynamics. The stone crab (Menippe spp.) is found throughout southwestern North Atlantic waters, from North Carolina through the Gulf of Mexico and the Central American Caribbean, including the Greater Antilles. Recent observations suggest that stone crabs have become better established on certain oyster reefs in North Carolina than in the early 1900s when they we first observed in NC. To assess the predatory impact of stone crabs on oysters, we (1) quantified stone crab densities on subtidal oyster reefs in Pamlico Sound, NC using scuba surveys, and (2) conducted laboratory predation experiments to assess the functional response of stone crabs to varying densities of oysters. We then (3) analyzed previously unpublished functional response data on another important oyster predator, the mud crab Panopeus herbstii. Finally, we (4) compared and contrasted potential predatory impacts of stone, mud and blue crabs (Callinectes sapidus). The functional response data and analyses for both stone crabs and mud crabs were consistent with a type II functional response. Mud crabs, on a m² basis, inflicted the highest proportional mortality on oysters over a 24 hour period, followed by stone and then blue crabs. Proportional mortality did not vary significantly with oyster size; however, relatively small and large oysters were consumed disproportionately less than medium-sized oysters, likely due to the mechanical inability of stone crabs to handle small oysters, and the inability to crush large oysters. Although stone crabs appear to be established in Pamlico Sound at densities equivalent to densities in other systems such as the U.S. Florida Panhandle, their predatory activities on oysters are not expected to have as significant a negative impact on oyster populations compared to other resident predators such as mud crabs.     

The density and spatial arrangement of the invasive oyster Crassostrea gigas determines its impact on settlement of native oyster larvae

Understanding how the density and spatial arrangement of invaders is critical to developing management strategies of pest species. The Pacific oyster, Crassostrea gigas, has been translocated around the world for aquaculture and in many instances has established wild populations. Relative to other species of bivalve, it displays rapid suspension feeding, which may cause mortality of pelagic invertebrate larvae. We compared the effect on settlement of Sydney rock oyster, Saccostrea glomerata, larvae of manipulating the spatial arrangement and density of native S. glomerata, and non‐native C. gigas. We hypothesized that while manipulations of dead oysters would reveal the same positive relationship between attachment surface area and S. glomerata settlement between the two species, manipulations of live oysters would reveal differing density‐dependent effects between the native and non‐native oyster. In the field, whether oysters were live or dead, more larvae settled on C. gigas than S. glomerata when substrate was arranged in monospecific clumps. When, however, the two species were interspersed, there were no differences in larval settlement between them. By contrast, in aquaria simulating a higher effective oyster density, more larvae settled on live S. glomerata than C. gigas. When C. gigas was prevented from suspension feeding, settlement of larvae on C. gigas was enhanced. By contrast, settlement was similar between the two species when dead. While the presently low densities of the invasive oyster C. gigas may enhance S. glomerata larval settlement in east Australian estuaries, future increases in densities could produce negative impacts on native oyster settlement. Synthesis and applications: Our study has shown that both the spatial arrangement and density of invaders can influence their impact. Hence, management strategies aimed at preventing invasive populations reaching damaging sizes should not only consider the threshold density at which impacts exceed some acceptable limit, but also how patch formation modifies this.     

Characterisation of microsatellite loci in the habitat-forming kelp, Ecklonia radiata (Phaeophyceae, Laminariales)

Habitat forming algae play an important role in the ecology of temperate reefs worldwide. Despite this, our understanding of levels of gene flow within and among populations of algae is largely limited to studies on intertidal species; we know comparatively little about important habitat-forming subtidal algae. Here, we develop eight polymorphic microsatellite markers for the characterisation of population genetic diversity and structure in the subtidal kelp, Ecklonia radiata. This large macroalga is the most abundant habitat-forming kelp on the subtidal rocky reefs of temperate Australia and New Zealand where it forms extensive forests that support an astounding diversity of associated taxa.     

Natural cultch type influences habitat preference and predation,but not survival,in reef‐associated species

A shared origin with fresh and dredged cultch and availability via mining have made fossil cultch a commonly used reef restoration substrate. However, important differences in shape and size between whole‐shell cultch and fossil cultch may impact the complexity of reefs constructed from these materials. To determine if these differences may impact the development of restored reefs, we quantified the interstitial space each cultch type provides and constructed reef mesocosms to measure (1) the immediate effects of exposure to each cultch type on mortality of blue crab (Callinectes sapidus) and pink shrimp (Farfantepenaeus duorarum); (2) the tendency of crab, shrimp, and Florida crown conch (Melongena corona) to be found on habitats composed of each substrate type and their position within each in split‐substrate mesocosms; and (3) the influence of cultch type on predation of Eastern oysters (Crassostrea virginica) by crabs and conch. Aggregation of fossil cultch contains more shells and provides less interstitial space than an equivalent volume of whole‐shell cultch. Although immediate mortality following deployment was low and did not differ among cultch types, we found that all species were more likely to be found on fresh cultch over fossil cultch in choice experiments and used each habitat type differently. Cultch type also impacted the size of oysters consumed by crabs in short‐term feeding trials. The structure and traits of habitats created by various materials should be added to the growing list of issues considered when natural communities are to be restored in oyster reefs and other environments.     

Experimental effects of kelp canopies on subtidal coralline algae

Plants are often grouped as canopy species or understorey species because it is thought that that these sets of taxa interact in predictable ways. Mensurative experiments in southern Australia demonstrated that the percentage cover of encrusting coralline algae was greater, and articulated (branching) coralline algae less, on boulders under a canopy of dense kelp (>7 plants per m²), Ecklonia radiata, than on boulders without kelp. Experimental clearances of kelp and reciprocal transplants of boulders between patches of E. radiata and patches without kelp showed that canopies maintained and facilitated the growth of encrusting coralline algae and reduced the cover of articulated coralline algae. Potential artefacts associated with clearing kelp and transplanting boulders were not detected when tested with a series of translocation controls. These results reject the model that the co‐occurrence of E. radiata and encrusting corallines is just an assemblage of plants caused by spatial and temporal coincidence. Instead, they support the model that kelp facilitates the growth and survival of understorey algae.     

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.     

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.     

Spatial variation in juvenile and adult Ecklonia radiata (Laminariales) sporophytes

Juvenile and adult sporophytes of Ecklonia radiata were counted on in-shore and off-shore reefs at Marmion (Perth, Western Australia), across spatial scales of kilometres, 10s of metres and metres. The position on in-shore reefs vs. off-shore reefs did not influence recruit and adult kelp density. There was considerable site-to-site (kilometres) variation in densities of recruits but not adults. The majority of variation in both recruit and adult densities was found between quadrats separated by a few metres. There was no relationship between abundance of recruits and abundance of adult sporophytes at any of the spatial levels of investigation, suggesting a decoupling of recruitment and canopy processes. The results emphasise the patchy nature of kelp canopy distribution and suggest that relatively small-scale processes, acting on scales of a few metres, are of primary importance in generating heterogeneity in the canopy of these kelps.     

Introduction,establishment and expansion of the Pacific oyster <Emphasis Type="Italic">Crassostrea gigas</Emphasis> in the Oosterschelde (SW Netherlands)

The Pacific oyster Crassostrea gigas was first introduced as an exotic species by oyster farmers in 1964 in the Oosterschelde estuary (SW Netherlands). The initial phase is not well documented but first natural spatfall was recorded in 1975. Excessive spatfall occurred in 1976 and this is considered the start of the expansion phase of the wild oysters. Oyster beds in intertidal and subtidal areas of the Oosterschelde estuary have been growing since. The development in the intertidal area has been reconstructed by using aerial photography, validated by ground truth in 2000–2002. In the subtidal areas extensive oyster beds have been detected by using side scan sonar; on hard substrates along the dikes coverage with oysters up to 90% locally has been recorded by scuba diving surveys. Expansion has also occurred into adjacent water bodies including the Wadden Sea. By forming resistant reefs the oysters induce structural changes in the ecosystem. It is concluded that bed area is still expanding while decrease of the fraction live animals may indicate adjustment of the stock size to the local conditions.     

Ecological interactions in the provision of habitat by urban development: whelks and engineering by oysters on artificial seawalls

Abstract Increases in human population cause increased urbanization of most habitats, including the shoreline. This has many consequences for coastal environments, in particular the trend for artificial structures, such as seawalls, to replace natural habitats. Seawalls and natural shores support many of the common intertidal species, but others important on rocky shores are absent from or rare on many seawalls. The whelk Morula marginalba Blainville is an abundant and important predator on rocky shores of south‐eastern Australia, but is infrequently recorded on artificial substrata. In Sydney Harbour, where the Sydney rock oyster (Saccostrea glomerata Gould) was locally abundant, densities of M. marginalba on some seawalls appeared to be similar to those on rocky shores and to be larger than where there were few oysters. We sampled densities and sizes of whelks in four habitats, predicting and corroborating that: (i) on seawalls with many oysters, there would be more whelks than on seawalls with few oysters; (ii) where there are many oysters, densities of whelks would be similar on seawalls and rocky shores; and (iii) whelks would be larger where oysters were abundant. Growth and survival of whelks were measured to test hypotheses from the observed differences in size and density. Survival was greater in habitats with many oysters, which could explain differences in density, but size‐specific differences in survival could not explain differences in size among habitats. On seawalls but not on rocky shores, slower growth could explain the smaller size of whelks where there were few oysters. Seawalls provide important habitat for M. marginalba, but only via their indirect effects, mediated by oysters. These interactions cannot be predicted from those on natural rocky shores. Predicting how developed areas provide suitable habitat, either in management of conservation or in assessments of potential impacts clearly depends on understanding the roles of biogenic habitats.     

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.     

A comparison of the predatory impacts of an invasive and native crab species using a functional response approach

The European green crab (Carcinus maenas) is invasive on the West coast of North America, but the ecological consequences of this invasion remain poorly understood. Comparative functional response analysis has arisen as a method of elucidating ecological consequences of invasive species by comparing the impact of these species to native analogues. Through comparative functional response experiments of green crabs and native red rock crabs (Cancer productus) we found that green crab predation increased asymptotically (Type II functional response) when fed increasing densities of Pacific oysters (Magallana gigas), while red rock crab predation displayed a sigmoidal (Type III) response. At high oyster densities red rock crabs consume more Pacific oysters than green crabs do, due to their reduced handling time, though green crabs consume more Pacific oysters relative to their size than red rock crabs. However, compared to red rock crabs, green crabs consume more oysters at low prey densities, which implies that they have a larger, potentially destabilizing impact on low densities of Pacific oysters. As green crabs continue to spread across the West coast of North America, Pacific oysters will face increased predation pressure. Our results show the advantage of using functional response analysis to compare density dependent predation between an invasive species and a native species to predict the ecological consequences of invasions.