s Abstract Biological invasions of marine habitats have been common, and many patterns emerge from the existing literature. In North America, we identify 298 nonindigenous species (NIS) of invertebrates and algae that are established in marine and estuarine waters, generating many "apparent patterns" of invasion: (a) The rate of reported invasions has increased exponentially over the past 200 years; (b) Most NIS are crustaceans and molluscs, while NIS in taxonomic groups dominated by small organisms are rare; (c) Most invasions have resulted from shipping; (d ) More NIS are present along the Pacific coast than the Atlantic and Gulf coasts; (e) Native and source regions of NIS differ among coasts, corresponding to trade patterns. The validity of these apparent patterns remains to be tested, because strong bias exists in the data. Overall, the emergent patterns reflect interactive effects of propagule supply, invasion resistance, and sampling bias. Understanding the relative contribution of each component remains a major challenge for invasion ecology and requires standardized, quantitative measures in space and time that we now lack.

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Allometric relationships of reproductive output were compared in 20 species from 7 families of brachyuran crabs from the east and west coasts of North America, using regression analysis of log reproductive parameters versus log body weight. Comparisons of crabs spanning 4 orders of magnitude in body weight indicated that female body size was the principal determinant of reproductive output: 95% of the variance in brood weight, 79% of the variance in the number of eggs per brood, 63% of the variance in annual brood weight, and 74% of the variance in annual fecundity were explained by body weight. Brood weight exhibited an isometric constraint to about 10% of body weight. Allometric limitations on space available for yolk accumulation in the body cavity appeared to be the main constraint on brood size. Ovum size increased significantly, but only slightly, with increasing body size. There was a significant trade-off between ovum size and the number of eggs per brood. There was no significant relationship between the number of broods per year and body size. The number of eggs per brood was significantly better than brood weight as a predictor of the number of broods produced per year by a species, indicating that demographic pressure on fecundity rather than energetic considerations is the primary selective mechanism influencing annual reproductive effort. Each of the 7 families of crabs exhibited trends toward distinct patterns for the suite of co-adapted reproductive traits. However, no interspecific reproductive patterns were apparent with respect to the variables of feeding type, salinity tolerance, habitat, and geographic range represented by the 20 species.

Invasions by non‐indigenous species (NIS) are recognized as important stressors of many communities throughout the world. Here, we evaluated available data on the role of NIS in marine and estuarine communities and their interactions with other anthropogenic stressors, using an intensive analysis of the Chesapeake Bay region as a case study. First, we reviewed the reported ecological impacts of 196 species that occur in tidal waters of the bay, including species that are known invaders as well as some that are cryptogenic (i.e., of uncertain origin). Second, we compared the impacts reported in and out of the bay region for the same 54 species of plants and fish from this group that regularly occur in the regionÂ's tidal waters. Third, we assessed the evidence for interaction in the distribution or performance of these 54 plant and fish species within the bay and other stressors. Of the 196 known and possible NIS, 39 (20%) were thought to have some significant impact on a resident population, community, habitat, or process within the bay region. However, quantitative data on impacts were found for only 12 of the 39, representing 31% of this group and 6% of all 196 species surveyed. The patterns of reported impacts in the bay for plants and fish were nearly identical: 29% were reported to have significant impacts, but quantitative impact data existed for only 7% (4/54) of these species. In contrast, 74% of the same species were reported to have significant impacts outside of the bay, and some quantitative impact data were found for 44% (24 /54) of them. Although it appears that 20% of the plant and fish species in our analysis may have significant impacts in the bay region based upon impacts measured elsewhere, we suggest that studies outside the region cannot reliably predict such impacts. We surmise that quantitative impact measures for individual bays or estuaries generally exist for <5% of the NIS present, and many of these measures are not particularly informative. Despite the increasing knowledge of marine invasions at many sites, it is evident that we understand little about the full extent and variety of the impacts they create singly and cumulatively. Given the multiple anthropogenic stressors that overlap with NIS in estuaries, we predict NIS‐stressor interactions play an important role in the pattern and impact of invasions.

Adult soft—shelled clams (Mya arenaria) persist at low densities in Chesapeake Bay sandy habitats despite intense predation by blue crabs (Callinectes sapidus). Clam persistence may be a consequence of variation in blue crab foraging rates as a function of clam density and sediment composition. In laboratory aquaria, we measured the functional responses (prey consumption per predator as a function of prey density) of large blue crabs to six densities of adult soft shelled clams buried at natural depths in two sediment types (mud and sand). Functional responses in sand and mud were differentiated statistically by analyses of (1) residuals and residual sums of squares of discrete and continuous—time models, and (2) the exponent b of a general functional response model. Crab predation rates were significantly higher in mud than sand, and functional responses differed significantly between these two substrates. Blue crabs displayed type III (sigmoid) density—dependent functional responses in sand, and type II (decelerating rise to an upper asymptote) inversely density—dependent responses in mud. Risk of mortality for clams decreased sharply in sand at low densities similar to those observed in the field near the end of the annual period of active predation. These observations (1) suggest that variable blue crab functional responses result in microhabitat—specific mortality rates of benthic prey, and (2) indicate that functional responses can differ significantly according to the physical properties of topographically simple habitats.

Introduced species frequently escape the natural enemies (predators, competitors, and parasites) that limit their distribution and abundance in the native range. This reduction in native predators, competitors, and parasites may result in ecological release in the introduced range. However, biological interactions also can limit the establishment and spread of nonnative populations. The extent to which such biotic resistance occurs is poorly resolved, especially for marine ecosystems. Here we test whether a native predator, the blue crab Callinectes sapidus, affects the abundance and geographic range of the introduced European green crab Carcinus maenas in eastern North America. Both crab species occur in shallow, soft‐sediment habitats of bays and estuaries, and their ranges overlap in eastern North America. First, we tested for a negative relationship in the abundances of the two species from trap samples across a 640‐km (5.78° latitude) coastal transect. Second, we estimated variation in predation pressure on tethered Carcinus maenas across latitude and as a function of Callinectes sapidus abundance. Third, we measured predation rates on Carcinus maenas by Callinectes sapidus in field and laboratory experiments. Our results support the hypothesis that the native predator Callinectes sapidus provides biotic resistance to invasion and prevents the southward spread and establishment of Carcinus maenas. Within and across bays, Carcinus maenas were significantly less abundant at sites and depths with Callinectes sapidus compared with areas lacking Callinectes sapidus. Moreover, no Carcinus maenas were found in Chesapeake Bay, where Callinectes sapidus were most abundant. Predation of tethered Carcinus maenas increased with Callinectes sapidus abundance. In laboratory and field experiments, Callinectes sapidus preyed readily on Carcinus maenas. Thus, we conclude the predation by Callinectes sapidus, alone or in combination with other factors, limits the abundance and geographic range of an invasive marine species.

The persistence of prey encountering intense predation varies by species, prey density, and habitat type; however, the collective impact of these factors has rarely been tested experimentally in natural marine systems. Using the thin‐shelled clams Mya arenaria and Macoma balthica as prey, and the main epibenthic predator of whole adult clams, the blue crab Callinectes sapidus, we conducted a series of experiments in Chesapeake Bay tributaries that (1) links field abundance and distribution of bivalve prey species with habitat‐specific mortality patterns; (2) represents the first comprehensive field test of species‐specific, habitat‐specific, and density‐dependent mortality for subtidal, soft‐bottom, deep‐burrowing prey; and (3) thereby enables development of a conceptual model to be used as a heuristic tool linking predator–prey dynamics, habitat type, and evolutionary defense tactics for marine benthos. In 15 years of field monitoring, Mya was more common in sand than mud habitats, and Macoma was widely distributed and at higher densities than Mya in mud and sand. In field experiments, mortality of both Mya and Macoma was density dependent in those habitats where the clams are common. The blue crab population in the field exhibited a type III “guild functional response” on Mya in sand, and on Macoma in both mud and sand. Mortality was lower in sand than mud for Mya, and similar in mud and sand for Macoma, correlating with the high abundances of Mya in sand and Macoma in sand and mud. The persistence of large juvenile and adult bivalves when confronted with intense predation derived substantially from a low‐density refuge from predation that varied in a species‐specific manner with habitat type, demonstrating the species‐specific importance of density and habitat to clam survival. We developed a conceptual model detailing the relative importance of behavior, morphology, habitat features, and the basic components of predator–prey interactions to the survival of bivalve molluscs. At one extreme are bivalve molluscs, such as oysters, that emphasize morphological refuges that increase the predator's handling time. At the other extreme are bivalves, such as Mya and Macoma, that reduce predator encounter rates. The model is intended to be used as a heuristic tool to develop testable hypotheses.

ABSTRACT. Abundances and size-frequency distributions of common epibenth~c flsh and crustaceans were compared among 3 depth zones (1-35, 35-70, 71-95 cm) of the Rhode River, a subestuary of Chesapeake Bay, USA. In the absence of submerged aquatic vegetation (SAV), inter-and intraspccific size segregation occurred by depth from May to October, 1989October, -1992. Small species (Palaemonetes pugjo, Crangon septernspjnosa, Fundulus heteroclitus, F majaljs, Rhithropanope~ls harrisii, Apeltes quadracus, Gobiosorna boscj) were most abundant at water depths <70 cm. Furthermore, the proportion of small individuals decreased significantly with depth for 7 of 8 species, with C. septemsp~nosa being the exception, exhibiting no size change with increasing depth. These distributional patterns were related to depth-dependent predalion risk. Large species (Callinectes sap~dus, Leiostomus xanthurus, and Micropogonias undulatus), known predators of some of the small species, were often most abundant in deep water (>70 cm). In field experiments, mortality of tethered P pugio (30 to 35 mm), small E heteroclitus (40 to 50 mm), and small C. sapjdus (30 to 70 mm) increased significantly with depth. Wc hypothesize that predation risk was size-dependent, creating the observed intra-and interspecific size differences among depth zones. For C. sep.temspinosa, burial may modify this size-dc>pen-dency and create the unusual absence of intraspecific size increase with depth. Historically, f ? pugio and Fundulus spp. (and other small species) were not restricted to shallow (<70 cm) waters and were abundant in deeper SAV beds, which provided a structural refuge from predators. Since the recent demise of SAV in Chesapeake Bay, our results indicate many small species have shifted their distributions and now utilize primarily shallow water as an alternate refuge habitat.