The effect of Southwestern Atlantic burrowing crabs on habitat use and foraging activity of migratory shorebirds

Southwestern Atlantic estuaries (Southern Brazil to Northern Patagonia, Argentina) are characterized by the presence of an intertidal burrowing crab Chasmagnathus granulata. This crab species is an important bioturbator which lives in large assemblages and excavates semi-permanent burrows that affect sediment characteristics. Our observations showed that distribution of the crabs in the Mar Chiquita coastal lagoon, Argentina (37°45′S, 57°26′W) affected habitat use and feeding success of migratory shorebirds. During the migratory season the two-banded plover (Charadrius falklandicus) foraged more frequently inside crab beds, and yellowlegs (Tringa flavipes and T. melanoleuca) fed more freqeuntly outside crab beds. Focal observations on the feeding behavior of the white-rumped sandpiper (Calidris fuscicollis) and the two-banded plover inside and outside crab beds showed that the plover was a visual searcher and captured more prey inside crab beds, and the white-rumped sandpiper was a tactile feeder. Although consumption rates (prey min^?1) did not differ between sites, their efficiency (prey probe^?1) inside crab beds was less. These differences were probably related to changes in sediment characteristics and prey behavior, which vary with crab activity. Burrowing crabs alter the suitability of intertidal habitats used by shorebirds in southwestern Atlantic estuaries. We believe that the same process could be occurring with other burrowing curstaceans such as thalassinidean shrimps in other estuaries of the world and could have important implications for management of flats for shorebirds. *** DIRECT SUPPORT *** A01BY090 00007     

Indirect effects of the intertidal burrowing crab<Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> in the habitat use of Argentina’s South West Atlantic salt marsh birds

The burrowing crab,Chasmagnathus granulatus, is the dominant benthic macroinvertebrate of southwest Atlantic salt marshes and tidal flats, having strong ecosystem engineering effects by direct and indirect effects on soil, vegetation, invertebrates, fishes, and birds. Vegetation structure is a main component for bird habitat selection, since greater habitat complexity generates higher niche diversity. This environmental complexity can be modified by species interactions or disturbance events (i.e., by crabs), in turn modifying the associated community. The bird species of salt marshes of the southwest Atlantic are highly dependent on these ecosystems. We assessed the effects of the burrowing crab on the structure of the cordgrass,Spartina densiflora, marshes, and how these changes affected the composition and diversity of birds. This study was conducted at the Mar Chiquita Coastal Lagoon, Argentina (37°40′S, 57°23′W).S. densiflora marshes were classified within three categories: marshes without burrowing crabs, marshes with low burrow densities, and marshes with high burrow densities. We made vegetation and bird surveys during spring beforeS. densiflora produce seeds and in summer when seeds are abundant. We also manipulated inflorescence (i.e., spike) density (a variable affected by crabs) to assess the effect on bird habitat use. The highest inflorescence density ofS. densiflora and highest bird diversity were recorded at low burrow densities. Cover and height ofS. densiflora were negatively correlated with burrow density. The number of bird species and the number of red-capped wren-spinetail,Spartonoica maluroides, were negatively related with cover and height ofS. densiflora. Experiments confirmed that granivorous species used areas with higher spike densities, a variable related to crab burrow density. Burrowing crabs directly and indirectly affect the cover, height, spike density, and morphologic characteristic of seeds ofS. densiflora marshes. These effects indirectly affect the bird community that uses these marshes, being negatively related toS. maluroides and total bird abundance, and positively related to bird diversity.     

The Intertidal Burrowing Crab <Emphasis Type="Italic">Neohelice</Emphasis> (=<Emphasis Type="Italic">Chasmagnathus</Emphasis>) <Emphasis Type="Italic">granulata</Emphasis> Positively Affects Foraging of Rodents in South Western Atlantic Salt Marshes

The role of positive and indirect interactions is often crucial in communities with intense abiotic stress such as salt marshes. The burrowing crab, Neohelice (=Chasmagnathus) granulata, is the dominant benthic macroinvertebrate of southwest Atlantic marshes (southern Brazil to Northern Argentinean Patagonia), having strong direct and indirect effects on marsh soil and, in consequence, on marsh vegetation and primary consumers. In this work, we investigate if this crab indirectly modifies habitat use by the granivorous rodents, Akodon azarae and Oligoryzomys flavescens, by increasing nutrient availability and thus enhancing seed production by the marsh plant Spartina densiflora. The study was conducted at the Mar Chiquita Coastal Lagoon, Argentina (37°32′ S). Rodent frequencies in S. densiflora were positively correlated with crab densities throughout the low and middle marsh. Additionally, the highest quality of S. densiflora and inflorescence density was recorded at the highest crab densities. Experimental manipulation of crab densities shows that N. granulata indirectly enhances the performance of S. densiflora (e.g., decreased fiber content and C/N ratios) and increases density of seeds. Moreover, N. granulata also facilitates S. densiflora seed availability to rodents by concentrating them in sediment mound at their burrows entrances. Experimental rodent exclusions showed that rodent species used S. densiflora seeds, a variable positively related to crab burrow density. Thus, our results show that N. granulata drives the granivorous rodent distribution and the intensity of seeds–rodent interaction trough facilitative and indirect interactions in marsh community.     

Predation on the southwestern Atlantic fiddler crab (Uca uruguayensis) by migratory shorebirds (pluvialis dominica, P. squatarola, arenaria interpres, and numenius phaeopus)

In Bahia Samborombon (SW Atlantic; 35°30′–36°22′S; 56°45′–57°23′W) migratory American golden plover (Pluvialis dominica), black-billed plover (P. squatarola), ruddy turnstone (Arenaria interpres), and whimbrel (Numenius phaeopus) forage on fiddler crabs (Uca uruguayensis). These shorebirds have distinct patterns of feeding behavior and capture different proportions of each sex and size. P. dominica made short runs, capturing mostly juvenile crabs, which are eaten whole. P. squatarola use a similar feeding strategy but capture females primarily. A. interpres walks continuously, capturing almost exclusively large male crabs (71%). N. phaeopus walk in the Uca patch and probe burrows by inserting the bill; they primarily capture females. The feeding rate of A. interpres is higher than that of P. dominica, P. squatarola, and N. phaeopus. Handling time shown by P. dominica and A. interpres was greater than the other two species for all categories of crabs. In all cases, handling time of male crabs was greater than those of either females or juveniles. All the evidences suggest that U. uruguayensis is an important food source for all these species and should be accounted in any conservation endeavor. *** DIRECT SUPPORT *** A01BY085 00006     

Effect of horseshoe crab spawning density on nest disturbance and exhumation of eggs: A simulation study

Because the Delaware Bay horseshoe crab (Limulus polyphemus) population is managed to provide for dependent species, such as migratory shorebirds, there is a need to understand the process of egg exhumation and to predict eggs available to foraging shorebirds. A simple spatial model was used to simulate horseshoe crab spawning that would occur on a typical Delaware Bay beach during spring tide cycles to quantify density-dependent nest disturbance. At least 20% of nests and eggs were disturbed for levels of spawning greater than one third of the average density in Delaware Bay during 2004. Nest disturbance increased approximately linearly as spawning density increased from one half to twice the 2004 level. As spawning density increased further, the percentage of eggs that were disturbed reached an asymptote of 70% for densities up to 10 times the density in 2004. Nest disturbance was heaviest in the mid beach zone. Nest disturbance precedes entrainment and begins the process of exhumation of eggs to surface sediments. Model predictions were combined with observations from egg surveys to estimate a snap-shot exhumation rate of 5–9% of disturbed eggs. Because an unknown quantity of eggs were exhumed and removed from the beach prior to the survey, cumulative exhumation rate was likely to have been higher than the snap-shot estimate. Because egg exhumation is density-dependent, in addition to managing for a high population size, identification and conservation of beaches where spawning horseshoe crabs concentrate in high densities (i.e., hot spots) are important steps toward providing a reliable food supply for migratory shorebirds.     

Effects of the intertidal burrowing crab<Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> on infaunal zonation patterns,tidal behavior,and risk of mortality

Rhythmic movements in response to tidal cycles are characteristic of infaunal inhabitant of intertidal soft-bottoms, allowing them to remain in the area with best living conditions. The effect of bioturbators as modifier of local environmental conditions and thus of gradients in intertidal habitats, has not been investigated yet. The Atlantic estuarine intertidal areas are dominated by the burrowing crabChasmagnathus granulatus that generates strong environmental heterogeneity by affecting the physical-chemical characteristics of the sediment. The comparison between intertidal areas with and without crab shows that sediments in the crab beds remain more humid, softer, and homogeneous across the intertidal and along the tidal cycle than areas without crabs. The densities of infauna were higher at high intertidal zones in crab beds than in similar areas without crabs. Infaunal organisms performed vertical movements into the sediment following the tidal cycle that were always of higher magnitude in habitats without crabs. Infaunal species tend to spend most of the time buried into the sediment in the crab bed. Migratory shorebirds use the Atlantic estuarine environments as stopover or wintering sites. They feed (mainly on polychaetes) in the low intertidal zones of both habitats (with and without crabs), but they also feed in the upper intertidal of the crab bed; polychaete per capita mortality rate is higher in the upper part of the crab bed. Environmental heterogeneity produced by crab disturbance has an effect on the infaunal behavior, risk of mortality, and the zonation pattern. This is another example of the ecosystem engineering ability of a burrowing intertidal species.     

The effect of migratory shorebirds on the benthic species of three southwestern Atlantic Argentinean estuaries

We experimentally evaluate the effect of migratory shorebirds on the benthic fauna of three southwestern Atlantic Argentinean stop-over and wintering sites: Bahía Samborombon (35°30′–36°22′S, 57°23′W), Mar Chiquita coastal lagoon (37°40′S, 57°26′W), and Bahía Blanca (38°48′–39°25′S, 50°–62°25′W). The experiments consisted of exclusion ceilings and controls (both 1 m²), with 10 replicates each and aligned at the same tidal level. During December 1994–May 1995 these experiments were done, twice in Bahía Samborombon, twice at two sites (Sotelo and Celpa) 4 km apart in Mar Chiquita coastal lagoon, and three times in Bahía Blanca. Three polychaete species (Laeonereis acuta, Nephtys fluviatilis, andHeteromastus similis) were found in similar densities in both areas of the Mar Chiquita lagoon.L. acuta andN. fluviatilis were affected in Sotelo, but there was no treatment effect in Celpa. In Sotelo the most abundant shorebirds were Hudsonian godwit (Limosa haemastica), American golden plover (Pluvialis dominica), and White-rumped sandpiper (Calidris fuscicollis), and in Celpa the most abundant shorebirds were White-rumped sandpiper, two-banded plower (Charadrius falklandicus), and lesser yellowlegs (Tringa flavipes). The largest polychaete densities were in the upper sediment layer (0–6 cm), which was also the most affected layer. Three polychaete species were found in Bahía Samborombon (L. acuta, H. similis, andNeanthes succinea), but onlyH. similis showed a treatment effect. The most abundant shorebirds in Samborombon were white-rumped sandpiper and two-banded plover. No effect was detected in Bahía Blanca, where the most abundant shorebirds were white-rumped sandpiper and American golden plover. In all cases, the species affected by shorebirds were the most abundant species. From the two sites of Mar Chiquita, there was a treatment effect only in Sotelo, which was also the area with higher shorebirds counts. However, much lower densities observed in Samborombon (similar to the unaffected area of Mar Chiquita) also produced a significant decrease in infaunal abundance. These evidences also suggest that just selection of study sites may produce different views of the interaction between shorebirds and benthic species.     

Horseshoe crabs (Limulus polyphemus) in an urban estuary (Jamaica Bay, New York) and the potential for ecological restoration

We assessed the suitability of intertidal habitats for spawning by horseshoe crabs (Limulus polyphemus) at 12 proposed restoration sites identified by the United States Army Corps of Engineers along the shore of Jamaica Bay, a highly developed estuary in New York City. Based on beach geomorphology, we chose to quantify horseshoe crab activity at five of the sites during the May–July 2000 breeding season. Horseshoe crabs spawned intensively on small patches of suitable sand within larger areas of eroding shoreline with bulkheads and rubble fill. Small areas of sand behind grounded barges at Brant Point and Dubos Point had densities of over 100,000 eggs m⁻², which was equal to or greater than the egg densities on longer, more natural appearing beaches at Spring Creek and Dead Horse Bay, or at a sand spit at Bayswater State Park. There were no significant differences in the percentage of Jamaica Bay horseshoe crab eggs that completed development when cultured using water from Jamaica Bay or lower Delaware Bay, a less polluted location. Only 1% of the embryos from Jamaica Bay exhibited developmental anomalies, a frequency comparable to a previously studied population from Delaware Bay. We suggest that the distribution and abundance of horseshoe crabs at our study areas in Jamaica Bay is presently limited by the availability of suitable shoreline for breeding, rather than by water quality. Restoration efforts that increase the amount of sandy beach in this urban estuary have a good likelihood of benefiting horseshoe crabs and providing additional value to migrating shorebirds that use horseshoe crab eggs as food.     

Seasonal movement and resource-use patterns of resident horseshoe crab (<Emphasis Type="Italic">Limulus polyphemus</Emphasis>) populations in a Maine,USA Estuary

Knowledge of resource-use and movement patterns is a missing component in the development of horseshoe crab (Limulus polyphemus) management strategies. Available evidence indicates the potential for a variety of possible migratory behaviors, but the lack of high-resolution, spatial-temporal data has hindered development of a year-round profile of ranging behavior. This need was addressed in the present study by using acoustic telemetry to track the movements of adult horseshoe crabs in two subembayments (Egypt and Hog Bays) of the Taunton Bay Estuary, Maine, from June 2003 to June 2005. Estimated mean total home range sizes were 64.1 and 61.4 ha for breeding crabs tagged in Egypt and Hog Bays, respectively. We observed no horseshoe crab dispersal to areas outside of the subembayments where they were tagged, so no mixing was observed between Egypt and Hog Bay individuals despite a < 4-km separation. Observed shifts in movement patterns, resource use (subtidal versus intertidal), and vagility facilitated a profile of seasonally partitioned horseshoe crab activity, which included late April to early May post-wintering, June–July breeding, August–September pre-wintering, and October–April wintering, where space usage represented about 10% of the mean total home range size. The apparent isolation of these resident populations implies a heightened vulnerability to overexploitation and large-scale habitat alteration that might be more easily sustained by larger, more vagile populations. This work underscores the need to apply horseshoe crab conservation, research, and management efforts at scales that are appropriate to the ranging patterns of crabs, which first requires application of high-resolution methods to identify those patterns.     

<Emphasis Type="Italic">Deepwater Horizon</Emphasis> Oil Spill Impacts on Salt Marsh Fiddler Crabs (<Emphasis Type="Italic">Uca</Emphasis> spp.)

The Deepwater Horizon oil spill was the largest marine oil spill in US waters to date and one of the largest worldwide. Impacts of this spill on salt marsh vegetation have been well documented, although impacts on marsh macroinvertebrates have received less attention. To examine impacts of the oil spill on an important marsh invertebrate and ecosystem engineer, we conducted a meta-analysis on fiddler crabs (Uca spp.) using published sources and newly available Natural Resources Damage Assessment (NRDA) and Gulf of Mexico Research Initiative (GoMRI) data. Fiddler crabs influence marsh ecosystem structure and function through their burrowing and feeding activities and are key prey for a number of marsh and estuarine predators. We tested the hypothesis that the spill affected fiddler crab burrow density (crab abundance), burrow diameter (crab size), and crab species composition. Averaged across multiple studies, sites, and years, our synthesis revealed a negative effect of oiling on all three metrics. Burrow densities were reduced by 39 % in oiled sites, with impacts and incomplete recovery observed over 2010–2014. Burrow diameters were reduced from 2010 to 2011, but appeared to have recovered by 2012. Fiddler crab species composition was altered through at least 2013 and only returned to reference conditions where marsh vegetation recovered, via restoration planting in one case. Given the spatial and temporal extent of data analyzed, this synthesis provides compelling evidence that the Deepwater Horizon spill suppressed populations of fiddler crabs in oiled marshes, likely affecting other ecosystem attributes, including marsh productivity, marsh soil characteristics, and associated predators.     

Evaluating salt marsh restoration in Delaware Bay: The response of blue crabs,<Emphasis Type="Italic">Callinectes sapidus</Emphasis>, at former salt hay farms

Marshes are important habitats for various life history stages of many fish and invertebrates. Much effort has been directed at restoring marshes, yet it is not clear how fish and invertebrates have responded to marsh restoration. The blue crab,Callinectes sapidus, uses marsh habitats during much of its benthic life. We investigated the response of blue crabs to marsh restoration by comparing crab abundance (catch per unit effort), mean size and size frequency distribution, sex ratio, and molt stages of crabs in recently restored marshes that were former salt hay farms to that of adjacent reference marshes with similar physical characteristics in the mesohaline portion of Delaware Bay. Field sampling occurred monthly (April–November) in 1997 and 1998 using replicate daytime otter trawls in large marsh creeks and weirs in smaller intertidal marsh creeks. Blue crabs were either equal or more abundant, the incidence of molting was in most months similar, and population sex ratios were indistinguishable in restored and reference marshes, suggesting that the restored areas attract crabs and support their growth. Site location had a greater effect on the sex ratio of crabs such that marshes closer to the mouth of the bay contained a higher percentage of adult female crabs. In each annual growing season (April–July), the monthly increase in crab size and, in some months (June–July), the incidence of molting at the restored sites was greater than the reference sites, suggesting that the restored sites may provide areas for enhanced growth of crabs. These results suggest that blue crabs have responded positively to restoration of former salt hay farms in the mesohaline portion of Delaware Bay.     

Trends in Abundance of Coastal Birds and Human Activity on a Texas Barrier Island Over Three Decades

Trends in abundance of the 28 most common species of coastal birds on Mustang Island, Texas, were assessed for a 29-year period (1979–2007) during which the study area experienced a substantial increase in human activity. Ten of the 28 species examined declined significantly (P < 0.05) over the study period. Six of these were larids (herring gull, Forster’s tern, royal tern, gull-billed tern, Caspian tern, black skimmer), which exhibited declines in mean abundance of 53% to 88%. Other significant species declines were the great blue heron (39% decline) and the shorebirds Wilson’s plover (63%), red knot (54%), and black-bellied plover (34%). Four species showed significant increases in local abundance. Mean number of people on the beach increased fivefold during the study period. This unique dataset allows for an extremely rare assessment of bird populations in direct relation to human disturbance and may have implications for managing coastal birds worldwide.     

Use of Multiple Chemical Tracers to Define Habitat Use of Indo-Pacific Mangrove Crab, <Emphasis Type="Italic">Scylla Serrata</Emphasis> (Decapoda: Portunidae)

The mangrove or mud crab, Scylla serrata, is an important component of mangrove fisheries throughout the Indo-Pacific. Understanding crab diets and habitat use should assist in managing these fisheries and could provide additional justification for conservation of the mangrove ecosystem itself. We used multiple chemical tracers to test whether crab movements were restricted to local mangrove forests, or extended to include adjacent seagrass beds and reef flats. We sampled three mangrove forests on the island of Kosrae in the Federated States of Micronesia at Lelu Harbor, Okat River, and Utwe tidal channel. Samples of S. serrata and likely food sources were analyzed for stable carbon (δ¹³C), nitrogen (δ¹⁵N), and sulfur (δ³⁴S) isotopes. Scylla serrata tissues also were analyzed for phosphorus (P), cations (K, Ca, Mg, Na), and trace elements (Mn, Fe, Cu, Zn, and B). Discriminant analysis indicated that at least 87% of the crabs remain in each site as distinct populations. Crab stable isotope values indicated potential differences in habitat use within estuaries. Values for δ¹³C and δ³⁴S in crabs from Okat and Utwe were low and similar to values expected from animals feeding within mangrove forests, e.g., feeding on infauna that had average δ¹³C values near −26.5‰. In contrast, crabs from Lelu had higher δ¹³C and δ³⁴S values, with average values of −21.8 and 7.8‰, respectively. These higher isotope values are consistent with increased crab foraging on reef flats and seagrasses. Given that S. serrata have been observed feeding on adjacent reef and seagrass environments on Kosrae, it is likely that they move in and out of the mangroves for feeding. Isotope mixing model results support these conclusions, with the greatest mangrove ecosystem contribution to S. serrata diet occurring in the largest mangrove forests. Conserving larger island mangrove forests (> 1 km deep) appears to support crab foraging activities.     

Organochlorine contaminants in a coastal lagoon in Argentina: Analysis of sediment, crabs, and cordgrass from two different habitats

Burrowing crabs (Chasmagnathus granulata), sediment collected from inside and outside crab burrows as well as outside the crab bed, and cordgrass (Spartina densiflora) were collected from intertidal mudflat and cordgrass marsh habitats and analyzed for concentrations of polychlorinated biphenyl (PCB) congeners and organochlorine (OC) compounds to test the hypothesis that there are differences in the distribution and bioaccumulation of OC contaminants in coastal lagoon habitats in Argentina. PCB concentrations were relatively low, although the penta- and hexachlorobiphenyl detected in sediments and biota indicated that there had been direct inputs of PCBs into the ecosystem. Heptachlor epoxide, dieldrin, endosulfan sulphate, chlordane compounds, DDT and metabolites, and hexachlorocyclohexanes (HCHs) were the major pesticides detected in sediment and biota samples. When lipid-normalized concentration data for all OC pesticides in crabs were summed together (ΣOCC), there were higher concentrations in crabs from the cordgrass habitat in comparison to crabs from the mudflat. In sediment samples, there were no significant differences in percent organic matter of marsh and mudflat sediments, but the concentrations of ΣOCC normalized to organic carbon were higher in the sediments collected in the cordgrass marsh. Samples of rhizomes and roots from the cordgrass contained high concentrations of OC compounds and it was estimated that 2.4 kg of heptachlor epoxide, the most abundant OC pesticide, may be present in the total cordgrass root biomass in Mar Chiquita lagoon. These data indicated that the cordgrass in coastal lagoon environments is an important factor in determining the distribution of persistent contaminants, and that a significant portion of the total burden of these hydrophobic compounds may be deposited in cordgrass biomass.     

Magnitude of harvest of Atlantic horseshoe crabs,<Emphasis Type="Italic">Limulus polyphemus</Emphasis>, in Pleasant Bay,Massachusetts

Atlantic horseshoe crabs,Limulus polyphemus, are currently harvested for biomedical, scientific, and bait purposes. In recent years, changes in population abundance and magnitude of harvesting have raised concerns about the status of this resource. We found horseshoe crab harvest in Pleasant Bay, Massachusetts, was selective by sex and size. Biomedical harvest preferred larger individuals and females, the scientific harvest preferred smaller individuals and males, and the bait harvest preferred females. Total 2001 harvest for all purposes accounted for the mortality of ∼1–2% the adult population. Biomedical harvest accounted for the greatest loss of horseshoe crabs from Pleasant Bay, ∼1–1.6% of the total population. Although biomedical harvest had the lowest associated mortality rate (∼10–15%), many more crabs were harvested from Pleasant Bay for biomedical purposes than for other uses. The scientific harvest accounted for the mortality of ∼0.4% of the population, and bait harvest accounted for the smallest mortality at ∼0.06% of the population. Harvest mortality rate was estimated to be lower in Pleasant Bay than in other Cape Cod areas and may be lower than natural mortality in the population. This study is the first qualitative investigation of commercial harvest on horseshoe crab populations and emphasizes that harvest pressures on different populations need to be individually evaluated.     

Blue crab,<Emphasis Type="Italic">Callinectes sapidus</Emphasis>, response to the invasive common reed,<Emphasis Type="Italic">Phragmites australis</Emphasis>: Abundance,size, sex ratio,and molting frequency

Much effort has been directed recently at restoring marshes, by the removal of the invasive common reed,Phragmites australis, yet it is not clear how fish and invertebrates have responded either to the invasion ofPhragmites or to marsh restoration. The blue crab,Callinectes sapidus, uses marsh habitats during much of its benthic life. We investigated the response of blue crabs toPhragmites invasion and restoration efforts by comparing crab abundance (catch per unit effort), mean size and size frequency distribution, sex ratio, and molting of crabs in three physically similar areas differing in marsh vegetation;Spartina-dominated,Phragmites-dominated, and a treated area (Phragmites removed and now dominated bySpartina) in one marsh in the upper portion of Delaware Bay. Field sampling occurred monthly (April to November) from 1999 to 2001 using replicate daytime otter trawls in large marsh creeks. Crabs were categorized by carapace width into recruits (<30 mm), juveniles (30–115 mm), and adults (>115 mm). Juveniles dominated the system, representing 69.4% of all crabs. Similar monthly increases in mean size and molting patterns during the growing season (May–August) occurred inSpartina (natural and treated sites) andPhragmites sites suggesting that, subtidal habitats, used for molting, in these areas do not differ. More juveniles in the feeding molt stage (i.e., intermolt) than in other molt stages and more recruits predominantly in the feeding molt stage than adults were inSpartina, suggesting differences in the marsh surfaces used as feeding habitats withSpartina being preferred. Sex ratios of each life history stage were skewed towards males, but this was related to the low salinity of Alloway Creek, rather than marsh surface vegetation. Our results suggest that marsh surface vegetation influences the way blue crabs use marsh surface habitats, thus restoration efforts focusing on changing vegetation type may have a positive influence on blue crabs.     

Foraging by Two Estuarine Fishes, <Emphasis Type="Italic">Fundulus heteroclitus</Emphasis> and <Emphasis Type="Italic">Fundulus majalis</Emphasis>, on Juvenile Asian Shore Crabs (<Emphasis Type="Italic">Hemigrapsus sanguineus</Emphasis>) in Western Long Island Sound

The Asian shore crab, Hemigrapsus sanguineus, is a recent and particularly successful introduction to the east coast of the USA. Little research has been done on the utilization of Asian shore crabs for food by native species, a potential form of biocontrol. Over a 4-year period, we examined the gut contents of cogeners, Fundulus heteroclitus and Fundulus majalis, collected from two embayments in western Long Island Sound for the presence of juvenile H. sanguineus. Frequency (percent) of occurrence of food items in the guts of both species varied over year and study site. Asian shore crabs were consumed more often by F. heteroclitus than by F. majalis, but predation pressure by both species was low. Only 13% of F. heteroclitus and 7.7% of F. majalis found with food in their guts had ingested Asian shore crab remains. Of those, 1/3 had consumed whole crabs; the rest had only autotomized appendages in their guts. The mean carapace width of juvenile Asian shore crabs ingested by F. heteroclitus was 3.59 ± 2.22 mm (N = 33). Results of our study on killifish predation support the hypothesis that H. sanguienus abundance is partly explained by reduced impact of native predators (i.e., the “enemy release hypothesis”). Predation pressure of other potential enemies on both planktonic and benthic stages of the Asian shore crab must be investigated, however, to understand the full impact of predation on H. sanguineus population dynamics.     

Decline in size of male blue crabs (Callinectes sapidus) from 1968 to 2000 near Calvert Cliffs, Maryland

Maryland commercial landings of the blue crab (Callinectes sapidus) and catch per unit effort (CPUE) have remained fairly stable over the past 33 yr despite occasional large deviations from the long-term average. During this time, however, significant declines in the percent of legal male crabs and the mean size of legal males caught in fishery-independent surveys near Calvert Cliffs, Maryland have become apparent. Sublegal females and two of the three legal female classes (152–177 and >178 mm CW) showed no significant trends over this 33-yr period when examined by linear regression. Males showed significant trends for all size classes. Sublegal males increased from 24% of the male population during the first 5 yr of the study (1968–1972) to 71% during 1996–2000. All classes of legal males, however, exhibited downward trends. Males 127–151 mm CW decreased from 45% of the male population in the earliest period to 22% during the last 5 yr. Males 152–177 mm decreased from 27% during 1968–1972 to only 6% during 1996–2000, and males > 178 mm declined from 4% in the earliest period to 0.5% in the recent period. These size decreases for the most valuable portion of the blue crab population are further evidence of over-exploitation. The declining trends in male size indicate that growth overfishing is occurring as intense fishing pressure removes so many male crabs from the population as they reach legal size that few remain to molt to larger size. A 3-yr data set from the Patuxent River, where commercial use of crab pots is restricted and fishing pressure is lower, suggests that legal male crabs are able to attain larger size compared to an area where the pot fishery is intense. A recommendation could be made for reducing effort where the pot fishery is intense by means of time, gear, catch limits, and/or by increasing the minimum size of legal crabs to allow larger crabs to enter the fishery.     

Bioenergetics modeling to investigate habitat use by the nonindigenous crab,Carcinus maenas, in Willapa Bay, Washington

A bioenergetics model was developed and applied to questions of habitat use and migration behavior of nonindigenous European green crab (Carcinus maenas) in Willapa Bay, Washington, USA. The model was parameterized using existing data from published studies on the ecology and physiology ofC. maenas and allied brachyuran crabs., Simulations of the model were run describing four scenarios of habitat use and behavior during a 214-d simulation period (April–October) including crabs occupying mid littoral habitat, high littoral habitat, sublittoral habitat, and sublittoral habitat but undertaking intertidal migrations. Monthly trapping was done along an intertidal gradient in Willapa Bay to determine the actual distribution of crabs for the same time interval as the simulation period, and model results were compared to the observed pattern. Model estimates suggest no intrinsic energetic incentive for crabs to occupy littoral habitats since metabolic costs were c. 6% higher for these individuals than their sublittoral counterparts. Crabs in the littoral simulations were also less efficient than sublittoral crabs at converting consumed energy into growth. Monthly trapping revealed thatC. maenas are found predominantly in mid littoral habitats of Willapa Bay and there is no evidence of resident sublittoral populations. The discrepancy intimates the significance of other factors, including interspecific interactions, that are not incorporated into the model but nonetheless increase metabolic demand. Agonistic encounters with native Dungeness crabs (Cancer magister) may be chief among these additional costs, andC. maenas may largely avoid interactions by remaining in littoral habitats neglected by native crabs, such as meadows of nonindigenous smooth cordgrass (Spartina alterniflora). AdultC. maenas in Willapa Bay may occupy tidal elevations that minimize such encounters, and metabolic costs, while simultaneously maximizing submersion time and foraging opportunities.     

Predation and Functional Responses of Carcinus maenas and Cancer magister in the Presence of the Introduced Cephalaspidean Philine orientalis

An increasing number of examples suggest that interactions among introduced species are ecologically important and relevant to the management of invaded systems. We investigated the potential for the introduced cephalaspidean sea slug Philine orientalis to interfere with the feeding of the introduced European green crab (Carcinus maenas) and the native Dungeness crab (Cancer magister). We observed co-occurrence of crab species and P. orientalis at field sites in Bodega Harbor and Tomales, San Pablo, and San Francisco Bays. In laboratory and field experiments, we determined whether crab feeding was suppressed by P. orientalis and the duration of this suppression for individual crabs. We also used foraging response models to explore changes in the feeding rate of crabs with varying densities of P. orientalis and small bivalve prey. We found that P. orientalis deterred predation by green and Dungeness crabs on small clams in laboratory feeding trials, but not in field experiments with green crabs and P. orientalis. Foraging models predicted that P. orientalis would only affect crab feeding in the field under specific conditions of crab, P. orientalis, and prey densities. These foraging models bridged an important gap between lab and field experiments and allowed us to predict how changes in species abundances at two trophic levels might alter the importance of crab suppression by P. orientalis.