Application of a Random Forest algorithm to predict spatial distribution of the potential yield of Ruditapes philippinarum in the Venice lagoon, Italy

We present a modelling framework that combines machine learning techniques and Geographic Information Systems to support the management of an important aquaculture species, Manila clam (Ruditapes philippinarum). We use the Venice lagoon (Italy), the first site in Europe for the production of R. philippinarum, to illustrate the potential of this modelling approach. To investigate the relationship between the yield of R. philippinarum and a set of environmental factors, we used a Random Forest (RF) algorithm. The RF model was tuned with a large data set (n = 1698) and validated by an independent data set (n = 841). Overall, the model provided good predictions of site-specific yields and the analysis of marginal effect of predictors showed substantial agreement among the modelled responses and available ecological knowledge for R. philippinarum. The most influent environmental factors for yield estimation were percentage of sand in the sediment, salinity, and water depth. Our results agree with findings from other North Adriatic lagoons. The application of the fitted RF model to continuous maps of all the environmental variables allowed estimates of the potential yield for the whole basin. Such a spatial representation enabled site-specific estimates of yield in different farming areas within the lagoon. We present a possible management application of our model by estimating the potential yield under the current farming distribution and comparing it to a proposed re-organization of the farming areas. Our analysis suggests a reduction of total yield is likely to result from the proposed re-organization.     

Population dynamics of naturalised Manila clams <Emphasis Type="Italic">Ruditapes philippinarum</Emphasis> in British coastal waters

The Manila clam Ruditapes philippinarum was introduced to Poole Harbour (lat 50°N) on the south coast of England in 1988 as a novel species for aquaculture. Contrary to expectations, this species naturalised. We report on individual growth patterns, recruitment, mortality and production within this population. On the intertidal mudflats the abundance of clams (>5 mm in length) varied seasonally between 18 and 56 individuals m⁻². There appear to be two recruitment events per year and there were 6 year classes in the population. A mid-summer decline in abundance was partly due to increased mortality but probably also a result of down-shore migration in response to high water temperatures and the development of anoxic conditions. A winter fishery removes c 75% of clams of fishable size (maximum shell length ≥40 mm) and c 20% of the annual production. The fishery depresses the maximum age and size attained by the clams but appears to be sustainable. Clam mortality due to factors other than fishing is highest in late-winter to early spring. The growth of the clams is intermediate in comparison with many published studies but remarkably good given their intertidal position. As on the coasts of the Adriatic Sea, where the clam is also non-native, the Manila clam has thrived in a shallow, eutrophic, lagoon-like system on the English coast. While the Poole Harbour population is currently Europe’s most northerly reported self-sustaining, naturalised population, given forecasts of increasing air and sea temperatures it might be expected that this species will eventually spread to more sites around the coasts of Northern Europe with associated economic and ecological consequences.     

Predation affects the susceptibility of hard clam Meretrix lusoria to Hg-stressed birnavirus

Predator–prey interaction in aquatic ecosystem is one of the simplest drivers affecting the species population dynamics. Predation controls are recognized as important aspects of ecosystem husbandry and management. In this paper we investigated how predation control cause an increase in host growth in the abundance of hard clam (Meretrix lusoria) populations subject to mercury (Hg)-stressed birnavirus. Here we linked predator–prey relationships with a bioenergetic matrix population model (MPM) associated with a susceptible–infectious–mortality (SIM) model based on a host–pathogen–predator framework to quantify the predator effects on population dynamics of disease in hard clam populations. Our results indicated that relative high predation rates could promote the hard clam abundances in relation to predators that selectively captured the infected hard clam, by which the disease transmission was suppressed. The results also demonstrated that predator-induced modifications in host behavior could have potential negative or positive effects on host growth depending on relative species density and resource dynamics. The most immediate implication of this study for the management of aquatic ecosystem is that, beyond the potential for causing a growth in abundance, predation might provoke greater predictability in aquatic ecosystem species populations and thereby increase the safety of ecosystem production from stochastic environmental events.     

Bioavailability of polycyclic aromatic hydrocarbons (PAHs) to short-neck clam (<Emphasis Type="Italic">Paphia undulata</Emphasis>) from sediment matrices in mudflat ecosystem of the west coast of Peninsular Malaysia

The bioaccumulation and bioavailability of polycyclic aromatic hydrocarbons (PAHs) were characterized in sediment and Paphia undulata (short-neck clam) from six mudflat areas in the west coasts of Peninsular Malaysia. The concentrations of total PAHs varied from 357.1 to 6257.1 and 179.9 ± 7.6 to 1657.5 ± 53.9 ng g ^?1 dry weight in sediment and short-neck clam samples, respectively. PAHs can be classified as moderate to very high level of pollution in sediments and moderate to high level of pollution in short-neck clams. The diagnostic ratios of individual PAHs and principal component analysis indicate both petrogenic and pyrogenic sources with significant dominance of pyrogenic source. The first PAHs biota-sediment accumulation factors and relative biota-sediment accumulation factors data for short-neck clam were obtained in this study, indicating a preferential accumulation of lower molecular weight PAHs. Evaluation of PAH levels in sediments and short-neck clams indicates that short-neck clam could be introduced as a good biomonitor in mudflats. The results also demonstrated that under environmental conditions, the sedimentary load of hydrocarbons appears to be one of the factors controlling their bioavailability to biota.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).     

Stable isotopes changes in the adductor muscle of diseased bivalve Ruditapes philippinarum

In this article, we show how a disease could bias stable isotope analyzes of trophic networks and propose a strategy in the choice of tissues to be analyzed. In the past few years, a new pathology (brown muscle disease or BMD) affecting the posterior adductor muscle of Ruditapes philippinarum has emerged in Arcachon Bay. BMD induces a necrosis of muscle tissues which become infused by conchiolin and hence calcified. As muscle of mollusks are often used for trophic food webs studies through stable isotopic analyzes, this work investigated the effect of BMD on carbon and nitrogen isotopic ratios of anterior and posterior adductor muscles of clams collected in February and August 2007. Infected clams displayed a lower condition index and a posterior adductor muscle δ¹³C enrichment of 1.2‰ in February and 0.7‰ in August. δ¹⁵N of posterior muscles was however not affected by the disease. Anterior muscle of diseased clams remained healthy and displayed the same isotopic signature as both posterior and anterior muscular tissues of healthy clam. Acidification significantly depleted δ¹³C in posterior muscles of infected clams, suggesting calcification, contrary to anterior muscles of infected clam and to both muscles of healthy clams, where no effect was observed. An X-ray diffractometry analysis confirmed the presence of CaCO₃ (aragonite). Trophic food web studies relying on stable isotope ratios should utilize only healthy animals or anterior adductor muscles when expertise in mollusk pathology is lacking.     

Clamming up: environmental forces diminish the perceptive ability of bivalve prey

The lethal and nonlethal impacts of predators in marine systems are often mediated via reciprocal detection of waterborne chemical signals between consumers and prey. Local flow environments can enhance or impair the chemoreception ability of consumers, but the effect of hydrodynamics on detection of predation risk by prey has not been investigated. Using clams as our model organism, we investigated two specific questions: (1) Can clams decrease their mortality by responding to predators? (2) Do fluid forces affect the ability of clams to detect approaching predators? Previous research has documented a decrease in clam feeding (pumping) in response to a neighboring predator. We determined the benefits of this behavior to survivorship by placing clams in the field with knobbed whelk or blue crab predators caged nearby and compared mortality between these clams and clams near a cage-only control. Significantly more clams survived in areas containing a caged predator, suggesting that predator-induced alterations in feeding reduce clam mortality in the field. We ascertained the effect of fluid forces on clam perception of predators in a laboratory flume by comparing the feeding (pumping) behavior of clams in response to crabs and whelks in flows of 3 and 11 cm/s. Clams pumped significantly less in the presence of predators, but their reaction to blue crabs diminished in the higher velocity flow, while their response to whelks remained constant in both flows. Thus, clam reactive distance to blue crabs was affected by fluid forces, but hydrodynamic effects on clam perceptive distance was predator specific. After predators were removed, clams exposed to whelks took significantly longer to resume feeding than those exposed to blue crabs. Our results suggest that prey perception of predators can be altered by physical forces. Prey detection of predators is the underlying mechanism for trait-mediated indirect interactions (TMIIs), and recent research has documented the importance of TMIIs to community structure. Since physical forces can influence prey perception, the prevalence of TMIIs in communities may, in part, be related to the sensory ability of prey, physical forces in the environment that impact sensory performance, and the type of predator detected.     

The behaviour of giant clams (Bivalvia: Cardiidae: Tridacninae)

Giant clams, the largest living bivalves, live in close association with coral reefs throughout the Indo-Pacific. These iconic invertebrates perform numerous important ecological roles as well as serve as flagship species—drawing attention to the ongoing destruction of coral reefs and their associated biodiversity. To date, no review of giant clams has focussed on their behaviour, yet this component of their autecology is critical to their life history and hence conservation. Almost 100 articles published between 1865 and 2014 include behavioural observations, and these have been collated and synthesised into five sections: spawning, locomotion, feeding, anti-predation, and stress responses. Even though the exact cues for spawning in the wild have yet to be elucidated, giant clams appear to display diel and lunar periodicities in reproduction, and for some species, peak breeding seasons have been established. Perhaps surprisingly, giant clams have considerable mobility, ranging from swimming and gliding as larvae to crawling in juveniles and adults. Chemotaxis and geotaxis have been established, but giant clams are not phototactic. At least one species exhibits clumping behaviour, which may enhance physical stabilisation, facilitate reproduction, or provide protection from predators. Giant clams undergo several shifts in their mode of acquiring nutrition; starting with a lecithotrophic and planktotrophic diet as larvae, switching to pedal feeding after metamorphosis followed by the transition to a dual mode of filter feeding and phototrophy once symbiosis with zooxanthellae (Symbiodinium spp.) is established. Because of their shell weight and/or byssal attachment, adult giant clams are unable to escape rapidly from threats using locomotion. Instead, they exhibit a suite of visually mediated anti-predation behaviours that include sudden contraction of the mantle, valve adduction, and squirting of water. Knowledge on the behaviour of giant clams will benefit conservation and restocking efforts and help fine-tune mariculture techniques. Understanding the repertoire of giant clam behaviours will also facilitate the prediction of threshold levels for sustainable exploitation as well as recovery rates of depleted clam populations.     

Mechanical clam dredging in Venice lagoon: ecosystem effects evaluated with a trophic mass-balance model

Harvesting of the invasive Manila clam, Tapes philippinarum, is the main exploitative activity in the Venice lagoon, but the mechanical dredges used in this free-access regime produce a considerable disturbance of the lagoon ecosystem. An ecosystem approach to study the complex effects of clam harvesting was implemented using a trophic mass-balance model. The trophic relations in the ecosystem were quantified with a mixed trophic impact analysis and further evaluated by considering different explanations for the " Tapes paradox", which consists of the apparent population enhancement of Manila clams by dredging and the apparent nutritional advantages that this species receives from re-suspended organic matter. The key-role played by this introduced species is highlighted by a network analysis that indicates a "wasp-waist control" of the system by Manila clams. The model constructed to characterise the present state of the Venice lagoon ecosystem is compared with models produced for a reconstructed past lagoon and a projected future lagoon. The future model was obtained by simulating the elimination of clam dredging in 10 years. The three different models were compared using thermodynamic and informational indices. Simulating the elimination of clam dredging produced a 33% increase in artisanal fishery catches, carried out by means of static gears, even with no change in fishing effort. These simulations also forecast an increase in the mean trophic level of the artisanal fishery catches as a positive effect of eliminating mechanical clam harvesting.     

Salinity and substratum effects on condition index of the bivalve Rangia cuneata

The condition index (CI) of the clam Rangia cuneata was measured monthly at 8 stations in 2 substrate types and 3 salinity regimes in the James River estuary, Virginia, USA. CI was higher in clams from sand bottoms than from mud, and decreased in both substrata in fresher water. Values were lowest in early spring and at a maximum in early autumn, with a pronounced autumn peak in the sand substratum. This pattern coincided with the annual salinity cycle. A field experiment showed that some factor associated with the water overlying a sand or mud substratum, rather than the nature of the substratum itself, was important in determining CI. The hypothesis is presented that this factor may be the higher suspended solids concentrations measured immediately over mud bottoms.Virginia Institute of Marine Science Contribution No. 782.     

A regional-scale sustainable development index: the case of Baja California Sur,Mexico

Sustainable development, as a multi-dimensional concept, is difficult to measure. Some efforts using indicators and indices have appeared in recent years, but most were developed on a national scale. Use of sustainability indicators has proven valuable for attaining better management of the environment by minimizing information gaps and maximizing community capabilities in terms of economic, social, environmental, and institutional sustainability dimensions. However, at least in the case of developing countries, the potential exists that national sustainability measures, based on national level indicators, may mask problems in sub-national zones with highly unsustainable conditions. This paper proposes a methodology to evaluate sustainable development at a local level, the use of which could be helpful in comparing different regions within a country or even among different countries. National sustainability indicators should result from a combination (whether additive or proportional) of regional sustainability indicators, as developed in this paper.     

Siphon nipping facilitates lethal predation in the clam Mesodesma mactroides (Reeve, 1854) (Mollusca: Bivalva)

In soft sediment marine communities, fishes frequently bite off extended siphons of buried clams; the consequential shortening of the siphon is known to reduce burial depth of the clams, secondarily increasing their vulnerability to lethal excavating predators. In this study, siphon nipping on the yellow clam, Mesodesma mactroides, was simulated by removing the top 6.6–30% of siphons. This caused a burrow reduction in 25–75%, respectively, compared to control individuals with intact siphons, in field and laboratory trials. To examine subsequent consequences of reduced burial depth, we exposed nipped and intact clams to potential predators in the laboratory simulating the observed natural clam abundance. Artificially nipped clams were consumed twice as much as control clams. The present results suggest that sympatric croppers contribute to the stock recovery failure by facilitation of lethal predation and that re-seeding to increase the local abundance of M. mactroides should be an essential aspect of conservation efforts in South America.     

UV-absorbing substances in zooxanthellate and azooxanthellate clams

The effects of UV-A and UV-B radiation on photosynthesis of zooxanthellae within the siphonal mantle of the giant clam, Tridacna crocea, and in isolation were studied. While UV-B irradiation (2.4 W m⁻², 20 min) completely suppressed photosynthesis of the isolated zooxanthellae, it had little effect on their photosynthetic ability if they were irradiated while within the siphonal mantle of the host tissue. Chemical analysis of the siphonal mantle of T. crocea showed the presence of significant amounts of mycosporine-like amino acids (MAAs), which absorb UV-A and -B light. However, no MAA was detected in the isolated zooxanthellae. MAAs were concentrated in the siphonal mantle and kidney tissues in comparison with other tissues. In the siphonal mantle, MAA concentrations were the highest in the outermost surface layer where most of the zooxanthella cells resided. This indicates that the zooxanthellae are protected from UV radiation by a screen of concentrated MAAs in the host clam. Aside from T. crocea, significant amounts of MAAs were found not only in other zooxanthellate clams, such as T. derasa, Hippopus hippopus, Colculum cardissa and Fragum unedo, but also in a closely related azooxanthellate clam, Vasticardium subrugosum. On the other hand, no MAA was detected in any of the zooxanthellae from these zooxanthellate clams. No MAA was detected in the tissues of a deep-sea bivalve, Calyptogena soyoae. Although MAAs seem to block strong UV radiation in the shallow-water clam, they are probably not essential for the clam's life in the dark. MAAs in shallow-water clams may be derived from food and accumulated in their tissues, especially in the siphonal mantle and kidney. Received: 29 November 1996 / Accepted: 13 January 1997     

Size-dependent photosynthetic performance in the giant clam <Emphasis Type="Italic">Tridacna maxima</Emphasis>, a mixotrophic marine bivalve

Giant clams form a symbiosis with photosynthetic algae of the genus Symbiodinium that reside in clam mantle tissue. The allometry of symbiont photosynthetic performance was investigated as a mechanism for the increasing percentage of giant clam carbon respiratory requirements provided by symbionts as clam size increases. Chlorophyll fluorescence measurements of symbionts of the giant clam Tridacna maxima were measured during experiments conducted in September of 2009 using specimens 0.5–200 g tissue wet weight (3–25 cm long), collected from waters around southern Taiwan (N 21°36′, E 120°47′) from July to August of 2009. Light-dependent decreases in effective quantum yield (∆F/F _m′) calculated as the noontime maximum excitation pressure over PSII (Q _m), relative electron transport rates (rETR), and dark-adapted maximum quantum yield (F _v/F _m) all varied as a quadratic function of clam size. Both Q _m and rETR increased as clam size increased up to ~10–50 g then decreased as clam size increased. F _v/F _m decreased as clam size increased up to ~5–50 g then increased as clam size increased. Chlorophyll fluorescence measurements of rETR were positively correlated with gross primary production measured during chamber incubations. Overall, symbionts of mid-sized clams ~5–50 g exhibited the highest light-dependent decreases in effective photosynthetic efficiencies, the highest relative electron transport rates, and the lowest maximum photosynthetic efficiencies, and symbiont photosynthetic performance is allometric with respect to host clam size.     

Analysis of the eastern Pacific yellowfin tuna fishery based on multiple management objectives

Vector optimization techniques were used to generate arbitrary segments of a policy frontier for a dynamic yellowfin tuna (Thunnus albacares) fishery model assuming fixed technology and considering four policy objectives: minimizing dolphin mortality, minimizing incidental catch (all species except dolphins), maximizing sustainable yield, and minimizing biological risk for the yellowfin tuna stock. Results show that along the policy frontier: (1) reducing incidental dolphin mortality increases the incidental catch of other species in a nonlinear way; (2) yield increases (subject to a biomass precautionary level) can only be obtained at the expense of higher levels of dolphin mortality and incidental catch; (3) biological risk increases as the level of tunas caught increases, but this increase depends on the type of fishery (longline fishing and three different modes of purse-seining: log-sets, dolphin-sets or school-sets) that dominates the fishing effort; (4) there is an indirect relationship between the dolphin mortality levels and those of biological risk; (5) there is a direct relationship between the incidental catch levels and biological risk. Catch obtained with dolphin-sets dominates the Pareto-optimal solutions with highest dolphin mortality levels but is associated with lower biological risk, whereas catch obtained with log-sets dominates in Pareto-optimal solutions with higher incidental catch and higher biological risk. In general, trade-offs or shadow prices among objectives are not linear, indicating that marginal costs vary along the policy frontier. Results of the trade-off analysis may provide useful information for decision-makers and other policy actors. Complete information about the preferences of the decision-makers regarding the objectives is necessary to recommend a specific management policy.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Deleterious effects of a nonPST bioactive compound(s) from <Emphasis Type="Italic">Alexandrium tamarense</Emphasis> on bivalve hemocytes

The known negative effects of shellfish toxin-producing dinoflagellates on feeding, burrowing and survival of some bivalve mollusks has prompted questions concerning whether they might also impair the internal defense system of affected bivalves and make them more susceptible to disease agents. The primary components of the cellular defense system are hemocytes. Many toxic dinoflagellates are too large to be ingested whole by hemocytes and would most likely be exposed to intracellular toxins only after the algae are consumed, broken down, and the water-soluble toxins, released. Therefore, we conducted a series of experiments in which hemocytes from two suspension-feeding bivalves—the Manila clam, Ruditapes philippinarum, and the softshell clam, Mya arenaria—were exposed in vitro to filtered extracts of one highly toxic paralytic shellfish toxin (PST)-producing and one nonPST-producing strain of Alexandrium tamarense (isolates PR18b, 76 ± 6 STXeq cell⁻¹ and CCMP115, with undetectable PST, respectively). We measured adherence and phagocytosis, two hemocyte attributes known to be inhibited by bacterial pathogens and other stressors. We found no measurable effect of a cell-free extract from a highly concentrated suspension of the PST-producing strain on hemocytes of either bivalve species. Instead, extract from the nonPST-producing strain had a consistent negative effect on both clams, resulting in significantly lower adherence and phagocytosis compared to strain PR18b and filtered seawater controls. The bioactive compound produced by strain CCMP115, which has yet to be characterized, may be similar to the PST-independent allelopathic compounds described for Alexandrium spp., which act on other plankters. These compounds and those produced by other harmful algae are known to cause immobilization, cellular deformation and lysis of co-occurring target organisms. Thus, nonPST producing Alexandrium spp., which do not cause paralysis and burrowing incapacitation of clams, may still produce a compound(s) that has negative effects not only on hemocytes, but on other molluscan cell types and their functions, as well.     

Population differences in nerve resistance to paralytic shellfish toxins in softshell clam, <Emphasis Type="Italic">Mya arenaria</Emphasis>, associated with sodium channel mutations

The softshell clam, Mya arenaria, is a commercially important bivalve with wide latitudinal distribution in North America. Populations of clams with a history of repeated exposure to toxic Alexandrium spp. have developed a natural resistance to the paralytic shellfish toxins (PSTs) produced by these algae. An association between PST resistance in individual clams and a single mutation in the saxitoxin (STX) binding region of the α-subunit of the voltage-gated sodium (Na⁺) channel gene was previously identified. Here we establish that more than one mutation associated with nerve resistance to STX occurred at this locus. Both cDNA from mRNA and genomic DNA sequences from individual clams are identical demonstrating that both alleles are expressed simultaneously. In addition, one resistant allele per individual is sufficient to confer neural resistance to STX even though heterozygous individuals show an intermediate level of resistance to STX in in vitro nerve trunk assays.     

Role of carbonic anhydrase in the supply of inorganic carbon to the giant clam—zooxanthellate symbiosis

The mechanism whereby inorganic carbon (C_i) is acquired by the symbiotic association between the giant clam (Tridacna derasa) and zooxanthellae (Symbiodinium sp.) has been investigated. C_i in the haemolymph of the clam is in equilibrium with the surrounding sea water. The photosynthesis rate exhibited by the intact clam varies as a function of the C_i concentration in the clam haemolymph. The gill tissue contains high carbonic anhydrase activity which may be important in adjusting the C_i equilibrium between haemolymph and sea water. Zooxanthellae (Symbiodinium sp.) isolated from the clam mantle prefer CO₂ to HCO ₃ ^- as a source of inorganic carbon. The zooxanthellae have low levels of carbonic anhydrase on the external surface of the cell; however, mantle extracts display high carbonic anhydrase activity. Carbonic anhydrase is absent from the mantle of aposymbiotic clams (T. gigas), indicating that this enzyme may be essential to the symbiosis. The enzyme is probably associated with the zooxanthellae tubes in the mantle. The results indicate that carbonic anhydrase plays an important role in the supply of carbon dioxide within the clam symbiosis.     

Invertebrate community responses to recreational clam digging

Marine reserves can help in maintaining biodiversity and potentially be useful as a fishery management tool by removing human-mediated impacts. Intertidal, soft-sediment habitats can often support robust recreational and commercial shellfish harvests, especially for clams; however, there is limited research on the effects of reserves in these habitats. In San Juan County, Washington, several reserves prohibit recreational clam digging. We examined the effects of these reserves on infaunal community composition through comparison with non-reserve beaches during a 6-week period. Clam abundance, overall species richness and total polychaete family richness were greater on reserve beaches compared to non-reserve beaches. Additionally, an experiment within a reserve demonstrated negative impacts of digging on non-target infauna. These effects probably resulted from local disruption and disturbance of the sediment habitat and not from increased post-digging predation, which was controlled. Intertidal reserves could play an important role in sustaining local and potentially regional biodiversity.