Biological consequences of ice rafting in a New England salt marsh community

Ice rafting of salt marsh peat is a recurrent phenomenon in north temperate regions. This process was simulated in a northern New England salt marsh to test several hypotheses concerning the effects of peat transport from high to low intertidal heights on the growth and mortality of key sessile organisms: the ribbed mussel Geukensia demissa (Dillwyn), the fucoid alga Fucus vesiculosus L. var. spiralis (Farlow) and the cordgrass Spartina alterniflora (Loisel.). Growth rates increased when Geukensia and Fucus were transported to the lower intertidal; however, Spartina died when similarly transported. Predation pressure (primarily from Carcinus maenus L.) on Geukensia was greater when it was rafted to the lower intertidal zone than in the upper intertidal habitat and was size specific; mussels >3.5cm reached a size-escape from crab predation.A winter survey of dislodged mussels revealed that 72% of the mussels collected were dead and 86% had been overgrown by large Fucus plants, >2.5 × the natural frequency of Fucus overgrowth (32%). In marsh habitats where hard substratum is rare, 91% of the Fucus were growing on Geukensia. A dislodgement experiment showed that a significantly greater percentage of Geukensia was dislodged after ice-out when Fucus was attached to the shell than those mussels without Fucus overgrowth. In the spring, a population survey conducted in the salt marsh examined densities, biomass and population structure of Geukensia, as well as densities, percent cover and biomass of Fucus. Values obtained in the foremarsh were compared to those from the peat islands recently rafted to the tidal flats. Both biomass and densities of Geukensia were similar in the two areas; however, the size-frequency distributions of the mussels were different. Since fewer large mussels, Fucus and Fucus-overgrown mussels were found on the newly transported peat islands, this pattern appears to reflect dislodgement of larger Geukensia by attached algae during ice transport. Two ice-related sources of mortality were identified for Geukensia: (1) Fucus overgrowth acted as a vector for mussel dislodgement and was an indirect source of mortality; and (2) ice crushing was a direct source of mortality for non-overgrown mussels.     

Testing a typology system of running waters for conservation planning in Hungary

The zebra mussel (Dreissena polymorpha) and its congener the quagga mussel (Dreissena rostriformis bugensis) are both invaders in freshwater, but have very different invasion histories, with zebra mussels attaining substantially faster rates of spread at virtually all spatial scales. However, in waterbodies where they co-occur, D. r. bugensis can displace D. polymorpha. To determine if the mechanisms for this displacement are associated with different survival and growth, we kept mussels in flow-through tanks for 289 days with two temperature regimes that mimicked the natural surface water (littoral zone) and hypolimnion conditions of Lake Erie. For the littoral zone regime, we used water directly from the surface of Lake Erie (range 4–25°C, average 11.9 ± 0.6°C). For the profundal zone treatment, Lake Erie surface water was chilled to about 6°C (range 5–8°C, average 6.2 ± 0.6°C) for the full duration of the experiment. For each of these temperature regimes, we used three replicate tanks with only zebra mussels present and three replicate tanks with only quagga mussels (150 ind./tank each), and three replicate tanks with both species (75 ind./tank of each species). Quagga mussels had higher survivorship and grew more than zebra mussels in all treatments. For both species, the size of the mussel entering the winter was critical for survivorship. Larger mussels had a higher survival over the winter in all treatments. For both species, there was a survivorship and growth tradeoff. In the warmer littoral zone treatment both species had higher growth, but lower survival than in the colder profundal zone treatment. Surprisingly, although quagga mussels outperformed zebra mussels, zebra mussel survivorship was better when they were faced with competition by quagga mussels than with just intraspecific competition. In addition, quagga mussels suffered size-specific mortality during the growing season only when facing interspecific competition with zebra mussels. Further experiments are needed to determine the possible mechanisms for these interspecific effects.     

Intraspecific life history variation in the threatened Louisiana pearlshell mussel, Margaritifera hembeli

1. Margaritifera hembeli is a threatened mussel limited to twenty-two headwater streams in the Red River drainage in central Louisiana, USA. This study evaluated intraspecific variation in density, growth, size and age structure and shell morphology among several isolated populations. This study also identified the host fish and considered the role that host fish distribution played in determining mussel recruitment. 2. Mussels were aggregated in beds and average densities differed among streams. However, maximum mussel densities in beds were similar in all streams; the observed maxima were among the largest for monospecific mussel beds in North America, often exceeding 300 individuals m^–2. 3. The maximum size reached by individuals differed among streams, but all size distributions were skewed towards larger individuals. A repeated measures analysis of tagged mussels in four populations, over a 2-year period, indicated 2-fold differences in growth rates among streams, and significant variation among years. Growth rates were not affected by local population density. Maximum ages reached, determined indirectly by comparing growth rates, varied from 45 to 75 years. A canonical discriminant analysis also revealed significant differences in shell morphology across populations. 4. Half of the populations showed evidence of recent recruitment, and these sites had fish assemblages dominated by the host fish Noturus phaeus (Taylor). Host fish abundance appeared more important than adult mussel density in explaining recruitment patterns. 5. Considerable intraspecific life history variation suggests that management strategies for this species should be stream-specific, with emphasis on ensuring long-term habitat stability.     

Comparative study of predatory responses in blue mussels (Mytilus edulis L.) produced in suspended long line cultures or collected from natural bottom mussel beds

Blue mussels (Mytilus edulis L.) are a valuable resource for commercial shellfish production and may also have uses as a tool in habitat improvement, because mussel beds can increase habitat diversity and complexity. A prerequisite for both commercial mussel production and habitat improvement is the availability of seed mussels collected with minimum impact on the benthic ecosystem. To examine whether mussels collected in suspended cultures can be used for bottom culture production and as tool in habitat improvement, the differences in predatory defence responses between suspended and bottom mussels exposed to the predatory shore crab (Carcinus maenas L.) were tested in laboratory experiments and in the field. Predatory defence responses (byssal attachment and aggregation) and morphological traits were tested in laboratory, while growth and mortality were examined in field experiments. Suspended mussels had an active response in relation to the predator by developing a significantly firmer attachment to the substrate and a closer aggregated structure. Bottom mussels had a passive strategy by having a thicker shell and larger relative size of the adductor muscle. In a field experiment mussels originated from suspended cultures had a higher length increment and lower mortality when compared to bottom mussels. It is concluded that suspended mussels potentially are an alternative resource to bottom culture and can be used in habitat improvement of mussel beds, but that the use of suspended mussels has to be tested further in large-scale field experiments.     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Predator‐induced morphological defences in two invasive dreissenid mussels: implications for species replacement

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Differential effects of the severe winter of 1995/96 on the intertidal bivalves Mytilus edulis, Cerastoderma edule and Mya arenaria in the Northern Wadden Sea

Intertidal mussel beds were severely damaged by scouring ice floes during the winter of 1995/96. Aerial surveys before and after the winter showed that more clusters of mussel beds vanished in a region with a higher areal share of tidal flats and a lower salinity, suggesting that the amount of ice present determined the magnitude of the disturbance on beds of Mytilus edulis. Nehls and Thiel [(1993) Neth J Sea Res 31:181–187] observed a strikingly similar spatial pattern of disturbances caused by severe storms in the Wadden Sea. Areas on mussel beds mechanically undisturbed by ice showed no reduced abundance and biomass of mussels, indicating that temperature alone was of little importance as a lethal factor. Conversely, Cerastoderma edule was strongly affected by low temperature. On average 80% died during the winter with extinctions up to 100% in the high tidal zone. At the lowest tidal level, surviving cockles were larger than those killed by the frost. A reinvestigation of sampled sites in autumn revealed that substantial further mortality had occurred during spring and summer which may constitute a time-lag effect of the preceding winter. There was no increased mortality in juvenile and adult Mya arenaria during the winter of 1995/96, confirming that this clam is a hard-winter species like Macoma balthica. Received in revised form: 7 May 2001 Electronic Publication     

Population structure of the ribbed mussel <Emphasis Type="Italic">Geukensia demissa</Emphasis> in salt marshes in the southern Gulf of St. Lawrence,Canada

The ribbed mussel, Geukensia demissa, is highly dependent on the cordgrass Spartina alterniflora for amelioration from environmental stress and substrate stabilization. Spartina alterniflora is a foundation species in marshes, and G. demissa is typically associated with cordgrass beds. Marshes in the southern Gulf of St. Lawrence are experiencing erosion and degradation, presumably as a result of increases in sea level, which increases salinity exposure and negatively impacts S. alterniflora. The population structure of the ribbed mussel, Geukensia demissa, was studied at nine sites in six estuaries in the southern Gulf of St. Lawrence in Nova Scotia, Canada, where marsh degradation is occurring. Mussel length was used as a proxy for age of G. demissa in three salt marsh zones characterized by density and elevation of Spartina alterniflora: (1) a lower zone in which the S. alterniflora was dead, but where the basal mat was coherent, (2) a zone of living, but low density S. alterniflora at the margin of the living marsh, and (3) a zone of dense S. alterniflora one to three meters back from the edge. Mussel length was significantly different across the three zones in seven of the nine sites. Mean length decreased as elevation increased, and small mussels (i.e., 1–3 cm) were absent at seven sites. The smallest mussels occurred in the dense S. alterniflora zone, higher in the marsh. Mussel length in the two western sites did not differ between zones, and small mussels (i.e., 1–3 cm) were present, but rare. The absence of small mussels in seven of the nine sites, and the size frequency distribution at remaining sites, suggests a lack of recent recruitment and a long-term threat to the survival of G. demissa. Salt marsh degradation and the death of S. alterniflora have negatively impacted G. demissa recruitment, and population decline is evident.     

The mechanics of predation by the shore crab,Carcinus maenas (L.), on the edible mussel,Mytilus edulis L.

Summary Mechanical aspects of predation by the shore crab, Carcinus maenas, on the edible mussel, Mytilus edulis, were examined. The shore crabs from the population studied utilized five distinct, largely size-related, mussel-opening techniques. Crushing the mussel umbone appeared the most successful opening method for medium-sized prey. Small mussels were crushed outright and large mussels could be opened by a slow, uneconomical, boring technique. The strengths of mussels, from an exposed shore, were tested under compression in four separate planes to determine the loads a crab would need to apply to crush the shells outright and the mechanical properties of mussels. Little inter-plane variability in compressive strength was observed, although intra-plane variability appeared high. The compressive strengths of mussels from a sheltered shore were found to be significantly higher than those from the exposed shore in the plane tested. A strain gauge was embedded in a mussel shell enabling the pattern and magnitude of forces produced by crab chelae in opening a mussel to be studied. The crab's chelae did not appear overwhelmingly strong when compared directly to the compressive strength of the crab's preferred mussel sizes. It is, therefore, postulated that crabs usually seek out and exploit weak spots in the umbone of mussels by trial and error, eventually breaking through the shell by a cumulative process of extending minute fractures in the shell substructure.     

Biological control of streptococcal infection in Nile tilapia Oreochromis niloticus (Linnaeus, 1758) using filter‐feeding bivalve mussel Pilsbryoconcha exilis (Lea, 1838)

Since bivalve mussels are able to graze heavily on bacteria, in this paper it is hypothesized that when mussels are cultured with fish, the filtering efficiency of the mussels will keep the bacterial population below a certain threshold and thus assist in reducing the risk of bacterial disease outbreaks. The ability of the filter‐feeding bivalve mussel Pilsbryoconcha exilis to control Streptococcus agalactiae was tested in a laboratory‐scale tilapia culture system. Juvenile Nile tilapia (Oreochromis niloticus), the bivalve mussel as well as the bacteria were cultured at different combinations using four treatments: treatment‐1: mussel and bacteria but no fish, treatment‐2: tilapia and mussel but no bacteria, treatment‐3: tilapia and bacteria but no mussel, and treatment‐4: tilapia, mussels, and bacteria. All treatments were run in three replicates; stocking rates were 10 tilapia juveniles; five mussels; and about 3.5 × 10⁵ colony forming units (CFU) ml^?1 of bacteria in 50‐L aquaria with 40‐L volume. The mussel reduced the bacterial population by 83.6–87.1% in a 3‐week period whereas in the absence of the mussel, the bacterial counts increased by 31.5%. Oresence of the mussel also resulted in significantly higher growth and lower mortality of tilapia juveniles than when the mussel was absent. The results of this experiment suggest that the freshwater mussel P. exilis could control the population of S. agalactiae in a laboratory‐scale tilapia culture system. Future studies should focus on the dynamic interactions among fish, mussels, and bacteria as well as on how input such as feed and other organic materials affect these interactions.     

Long-term demography of a zebra mussel (Dreissena polymorpha) population

1. We used long‐term data and a simulation model to investigate temporal fluctuations in zebra mussel populations, which govern the ecological and economic impacts of this pest species. 2. The size of the zebra mussel (Dreissena polymorpha) population in the Hudson River estuary fluctuated approximately 11‐fold across a 13‐year period, following a cycle with a 2–4 year period. 3. This cycling was caused by low recruitment during years of high adult population size, rapid somatic growth of settled animals, and adult survivorship of 50% per year. 4. Adult growth and body condition were weakly correlated with phytoplankton biomass. 5. The habitat distribution of the Hudson's population changed over the 13‐year period, with an increasing proportion of the population spreading onto soft sediments over time. The character of soft‐sediment habitats in the Hudson changed because of large amounts (mean = 34 g DM m^?2) of empty zebra mussel shells now in the sediments. 6. Simulation models show that zebra mussel populations can show a range of long‐term trajectories, depending on the balance between adult space limitation, larval food limitation, and disturbance. 7. Effective understanding and management of the effects of zebra mussels and other alien species depend on understanding of their long‐term demography, which may vary across ecosystems.     

Biotic resistance on the increase: native predators structure invasive zebra mussel populations

1. Abundant native predators, parasites and pathogens that switch to consuming a hyper‐successful exotic species may be able to control the invasive population. Native predators may, however, need time to adapt to feed effectively on an exotic resource. In this case, mortality on an exotic population from native predators could increase over time even without a numerical increase in the predator population. 2. We measured mortality of zebra mussels (Dreissena polymorpha) in the Hudson River both in controls open to predation and in exclosures that excluded large predators to estimate mortality of zebra mussels from large predators and other causes. 3. We found that predation by the blue crab (Callinectes sapidus), and perhaps other predators, causes high mortality on zebra mussels in the Hudson River estuary. This predation apparently led to increased mortality and altered population structure in the invader over time. 4. Long‐term data from the Hudson River suggest that components of the invaded ecosystem, like rotifers, are recovering through predator‐caused release from zebra mussel grazing. Increased mortality on hyper‐successful exotic populations over time may be a common phenomenon with both ecological and management implications.     

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.     

Turbidity-mediated interactions between invasive filter-feeding mussels and native bioturbating mayflies

1. Invasive dreissenid mussels are known to cause large ecosystem changes because of their high filter‐feeding capacity, while native bioturbators may interfere with the mussels filter feeding. In this experiment, we investigated indirect environmental interactions between invasive filter‐feeding dreissenid mussels (zebra and quagga mussels) and native recolonizing bioturbating hexagenid mayflies (Hexagenia) at two mussel densities and two Hexagenia densities in a 2‐month long laboratory experiment. 2. Mean turbidity increased with increasing density of Hexagenia and decreased with increasing density of mussels. Turbidity showed the fastest decline at the highest mussel density, and no decline or a lower rate of decline at the low mussel density, dependent on Hexagenia density. 3. Mussel growth decreased with increasing Hexagenia density at low but not at high mussel density. Moreover, growth of mussels decreased as a function of increased mean turbidity at low mussel density but not at high mussel density. Filtering activity at the highest mussel density increased after introduction of food at the lower two densities of Hexagenia, but was constantly high at the highest Hexagenia density. 4. There was no difference in emergence of Hexagenia among the treatments, but mortality of Hexagenia was higher in the presence of mussels than in their absence. 5. Our results indicate that interactions between dreissenids and hexagenids are mediated through the sediment, and depend on density of both dreissenids and hexagenids. As the natural densities of these animals vary considerably within lakes, their growth and survival because of indirect environmental interactions is expected to vary spatially.     

Climate effects on inter‐annual variation in growth of the freshwater mussel (Anodonta beringiana) in an Alaskan lake

1. Warming trends are evident in many parts of the globe but are especially marked at higher latitudes, with complex effects on the biota that include direct effects on growth potential and indirect effects through food webs. 2. Air temperatures have been increasing over the past 50 years in southwestern Alaska, affecting the growth and population dynamics of many organisms, including a variety of aquatic species such as the freshwater mussel Anodonta beringiana. 3. We collected freshwater mussels from Iliamna Lake, in the Bristol Bay region of Alaska, and measured their shells to examine climatic effects on growth patterns. 4. Linear mixed effects models and ordinary least square linear regressions revealed strong positive correlations between local air temperatures (especially in May, October and the summer months) and inter‐annual variation in mussel growth. Slower mussel growth was also significantly correlated with later date of ice break‐up, which was linked to air temperatures in late spring.     

Suppression of microzooplankton by zebra mussels: importance of mussel size

1. The zebra mussel (Dreissena polymorpha) is amongst the most recent species to invade the Great Lakes. We explored the suppressive capabilities of mussels 6–22-mm in size on Lake St Clair microzooplankton (< 240)μm) in laboratory experiments. 2. Absolute suppression of rotifers and Dreissena veliger larvae was proportional to mussel shell length for individuals larger than 10 mm; larger zooplankton, mainly copepod nauplii and Cladocera, were not affected. Mussel clearance rates on rotifers generally exceeded those on veligers, although rates for both increased with increasing mussel size. Rotifer-based clearance rates of large (22 mm) mussels approached published values for phytoplankton food. 3. Most zooplankton taxa, particularly rotifers, declined significantly in western Lake Erie during the late 1980s concomitant with the establishment and population growth of zebra mussels in the basin. Densities of some taxa subsequently increased, although rotifers and copepod nauplii densities remained suppressed through 1993. Available evidence indicates that direct suppression by Dreissena coupled with food limitation provides the most parsimonious explanation for these patterns.     

Investigation of the effects of current velocity on mussel feeding and mussel bed stability using an annular flume

An annular flume was used to measure the effect of increasing current velocity on mussel (Mytilus edulis) feeding rate and the stability of mussel beds sampled from the mouth of the Exe estuary (SW England). It was found that, in contrast to earlier flume studies, the feeding rates of mussels from open coast sites were unaffected by current velocities up to 0.8 m s^–1. Algal cell depletion in the water column above mussels was a function of current velocity, increasing with declining currents below 0.05 m s^–1. The erodability/stability of the mussel bed, measured in terms of critical erosion velocity, sediment mass eroded and mean erosion rate, was found to be a function of the nature of the substrate and the density of the mussels. Erosion of mussel beds on sandy substrate showed a non-linear relationship with mussel bed density. In comparison with the sand (0% mussel cover), sediment resuspension was about five and four times higher for 25% and 50% cover, respectively. This was due to the increased turbulence and scouring around the clumps of mussels in low-density parts of the bed, and this resulted in some mussels detaching from the bed. At ~100% mussel cover, the sandy bed was more protected by the dense surface layer of mussels, and none became detached during erosion due to the high number of byssal attachments between individuals. The sediment resuspension from the 100% mussel cover was about three times lower than the 0% cover. Erosion of the bed with 50% cover resulted in burial of a large proportion of the mussels, with a 6 cm increase in sediment level. However, the mussels returned to the surface and recovered in 1–2 days, due to a combination of migration upwards and substrate settlement. Channels on the edge of the main Exmouth mussel bed were characterised by a more stable substrate comprising pebbles and sand with varying mussel densities. At these sites, where mussels experience high current velocities on spring tides (up to 0.9 m s^–1), there was no difference between the erodability of pebble/sand substrate with 0% and 100% mussel cover. The sediment erosion was also lower than the 100% mussel cover on the sandy substrate, particularly at currents >0.4 m s^–1. Sampling of different parts of the mussel bed at Exmouth showed mussels at low densities were made up of smaller clumps with a lower mass ratio of mussels to attached substrate (pebbles/sand), thus providing a greater degree of anchorage. Electronic Publication     

Optimal foraging versus shared doom effects: interactive influence of mussel size and epibiosis on predator preference

In the western Baltic Sea, the highly competitive blue mussel Mytilus edulis tends to monopolize shallow water hard substrata. In many habitats, mussel dominance is mainly controlled by the generalist predator Carcinus maenas. These predator-prey interactions seem to be affected by mussel size (relative to crab size) and mussel epibionts.There is a clear relationship between prey size and predator size as suggested by the optimal foraging theory: Each crab size class preferentially preys on a certain mussel size class. Preferred prey size increases with crab size.Epibionts on Mytilus, however, influence this simple pattern of feeding preferences by crabs. When offered similarly sized mussels, crabs prefer Balanus-fouled mussels over clean mussels. There is, however, a hierarchy of factors: the influence of attractive epibiotic barnacles is weaker than the factor ‘mussel size’. Testing small mussels against large mussels, presence or absence of epibiotic barnacles does not significantly alter preferences caused by mussel size. Balanus enhanced crab predation on mussels in two ways: Additional food gain and, probably more important, improvement in handling of the prey. The latter effect is illustrated by the fact that artificial barnacle mimics increased crab predation on mussels to the same extent as do live barnacles.We conclude that crab predation preferences follows the optimal foraging model when prey belong to different size classes, whereas within size classes crab preferences is controlled by epibionts.     

Recruitment of shore crabsCarcinus maenas on tidal flats: Mussel clumps as an important refuge for juveniles

During the late summer and early fall, juvenile shore crabs (Carcinus maenas L.) occurred in high abundances in mussel clumps scattered on tidal flats of the Wadden Sea. Abundances were much lower on bare tidal flats without mussel clumps and decreased substantially from July to November, whereas numbers in mussel clumps remained high. Large crabs left the tidal flats in early fall, whereas juveniles undertook tidal migrations only in the late fall. In March very few shore crabs were found in the intertidal area. The size of juvenile shore crabs living between mussels did not increase significantly during fall. On the bare tidal flats surrounding the mussels, a size increase was observed. Mussel beds and mussel clumps serve as a spatial refuge for the early benthic phases of juvenile shore crabs. Between mussels they can hide effectively from their epibenthic predators. Juvenile shore crabs do not leave the intertidal area and the mussel habitats before their major predators have left the area. Mussel clumps scattered over the tidal flats may be a critical refuge for juvenile shore crabs settling on tidal flats. Intensified efforts in mussel culturing in the European Wadden Sea during recent decades may have caused an increased abundance of mussel clumps on tidal flats, thus enhancing habitat availability for some mussel-clump inhabitants.     

Population growth of a gorgonian coral: equilibrium and non-equilibrium sensitivity to changes in life history variables

Summary A size dependent model of population growth of the Caribbean gorgonian Plexaura A is developed based on observed rates of survival, growth and colony fragmentation at a site in the San Blas Islands, Panama. Sensitivity and elasticity analyses indicate that the fate of large colonies has the greatest effect on population growth. Variables which directly affect the generation of large colonies have the next greatest effect on population growth. These variables include the recruitment of large fragments, and the survivorship of colonies in the next smaller size class. Sexual reproduction has an extremely limited ability to affect population growth. Vegetative reproduction has a greater potential effect on growth rates. Environmental conditions regularly change the matrix of transition probabilities which predicts population growth. This keeps the population from approaching its stable size class distribution. Deviations from the stable size class distribution alter sensitivity and elasticity and in this case have the effect of increasing the importance of survivorship of the smallest colonies. Nonequilibrium conditions alter sensitivity analyses and it is important to assess whether populations are at equilibrium and to determine the effects of such deviations on the sensitivity analysis.