Gregory M. Ruiz

Global spread of microorganisms by ships

Commercial ships have spread many species around the world, but little is known of the extent and potential significance of ship-mediated transfer of microorganisms. Here we show that the global movement of ballast water by ships creates a long-distance dispersal mechanism for human pathogens and may be important in the worldwide distribution of microorganisms, as well as for the epidemiology of waterborne diseases affecting plants and animals.

THE IMPACTS OF A NONINDIGENOUS MARINE PREDATOR IN A CALIFORNIA BAY

Coastal marine ecosystems worldwide are being altered rapidly by the invasion of nonindigenous species. Unlike terrestrial and freshwater systems, the impacts of an invading species have never been quantified on multiple trophic levels for a marine food web. We measured the impact of the nonindigenous green crab, Carcinus maenas, on a coastal marine food web in central California and found that this predator exerted strong “top-down” control, significantly reducing the abundances of several of the 20 invertebrate species monitored over a 9-yr period. Densities of native clams, Nutricola tantilla and Nutricola confusa, and native shore crabs, Hemigrapsus oregonensis, showed 5-fold to 10-fold declines within 3 yr of the arrival of green crabs. Field and laboratory experiments indicated that green crab predation caused these declines. We also tested for indirect responses of invertebrates and vertebrates to green crab predation. There were significant increases in the abundances of two polychaete taxa, Lumbr...

Predicting the impact of introduced marine species : Lessons from the multiple invasions of the European green crab Carcinus maenas

We compared ecological characteristics of three spatially independent invasions of the European green crab Carcinus maenas to determine which characteristics were most consistent across invasions, and hence would be most predictable in future invasions. For invasions in western North America (WNA), eastern North America (ENA), and South Africa (SAF), we compared five characteristics: (1) habitat usage, (2) diet preferences, (3) size of individuals, (4) rate of range expansion, and (5) demonstrated and potential impacts. We found that two characteristics, diet preference and ecological impact were relatively similar across the three invasions. Diet preference was particularly consistent with the rank order of taxa being virtually identical at the three sites. In contrast, habitat usage, individual size, and rate of range expansion were more variable. Differences in habitat usage and size were particularly evident in the WNA invasion, where C. maenas have failed to colonize protected and exposed rocky shores used elsewhere and have grown much larger than at other sites. We suggest that the degree of similarity of these characteristics across invasions provides a valuable measure of how predictable they will be in future invasions.

BIOTIC RESISTANCE TO INVASION: NATIVE PREDATOR LIMITS ABUNDANCE AND DISTRIBUTION OF AN INTRODUCED CRAB

Introduced species frequently escape the natural enemies (predators, com- petitors, 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 in- troduced European green crab Carcinus maenasin 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 (5.788 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 maenasby Callinectes sapidusin field and laboratory experiments. Our results support the hypothesis that the native predator Callinectes sapidusprovides 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 sapiduscompared with areas lacking Callinectes sapidus. Moreover, no Carcinus maenaswere found in Chesapeake Bay, where Callinectes sapidus were most abundant. Predation of tethered Carcinus maenasincreased with Callinectes sapidusabun- dance. 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.

Spread and potential impact of the recently introduced European green crab, Carcinus maenas, in central California

Our study examines the potential impact of the European green crab Carcinus maenas on communities of coastal embayments of western North America. We document the current distribution and range expansion of this species beyond San Francisco Bay, where C. maenas first became established along this coast in 1989–1990, and we test the effect of C. maenas predation on different species and sizes of infaunal invertebrates in field and laboratory experiments. In our samples from eight coastal locations in central California collected between June 1993 and May 1994, we found no green crabs at the two closest embayments south of San Francisco Bay and found the crabs in all four embayments sampled within 120 km north of San Francisco Bay, up to and including Bodega Harbor. C. maenas was not present in samples from sites farther north. This northward range expansion is apparently the result of larval recruitment by a single cohort, corresponding to the predominant northern transport of surface waters and the approximate distance water moves during larval green crab development. At Bodega Harbor, the current northern range limit, the C. maenas population is now well established and reproducing. Females and males became sexually mature within their first year at ≃40 mm carapace width, molting approximately monthly from summer through fall, and females were ovigerous in late fall of their first year at ≃50 mm. We expect larvae from this population to recruit locally and to the north, promoting episodic range extensions as new populations are established and reproduce. Enclosure experiments conducted during the summer of 1993 at the intertidal sandflats of Bodega Harbor showed that C. maenas significantly reduced densities of the most abundant taxa, including the bivalves Transennella confusa and T. tantilla, the cumacean Cumella vulgaris, and the amphipod Corophium sp. Furthermore, Carcinus maenas selectively removed larger (>3 mm) rather than smaller (<1 mm) Transennella spp. in both field and laboratory experiments. Based on the available data from this and other studies of green crabs, and our 10 yr study of community dynamics at Bodega Harbor, we predict C. maenas will significantly alter community structure, ecological interactions, and evolutionary processes in embayments of western North America.

Cryptic invasions of the crab Carcinus detected by molecular phylogeography.

Coastal marine ecosystems world‐wide are threatened by invasions of nonindigenous species. The ubiquity of marine sibling species identifiable only by genetic analysis suggests that many invasions are cryptic and therefore undetected, causing an underestimation of the actual number and impacts of invading species. We test this hypothesis with European crabs in the genus Carcinus that have invaded five regions globally. Partial 16S ribosomal RNA gene sequences confirm sibling species status of morphologically similar Atlantic C. maenas and Mediterranean C. aestuarii. Based on 16S rRNA haplotypes, crabs from California, New England and Tasmania were all C. maenas. However, we report the cryptic multiple invasion of both species in Japan and South Africa, where only C. aestuarii and C. maenas, respectively, were previously recognized.

Coarse woody debris as a refuge from predation in aquatic communities

This study demonstrates experimentally that coarse woody debris (CWD) can provide refuge from predation in aquatic habitats. In the Rhode River subestuary of Chesapeake Bay, Maryland, (USA), we (1) measured the abundance of CWD, (2) examined the utilization of CWD by mobile epibenthic fish and crustaceans, and (3) tested experimentally the value of CWD as a refuge from predation. CWD was the dominant above-bottom physical structure in shallow water, ranging in size from small branches (<2 cm diameter) to fallen trees (>50 cm diameter). In response to experimental additions of CWD, densities of common epibenthic cpecies (Callinectes sapidus, Fundulus heteroclitus, Fundulus majalis, Gobiosoma bosc, Gobiesox strumosus, Palaemonetes pugio, and Rithropanopeus harrisii) increased significantly compared to control sites without CWD. In laboratory experiments, grass shrimp (P. pugio) responded to predatory fish (F. heteroclitus and Micropogonias undulatus) by utilizing shelter at CWD more frequently than in absence of fish. Access to CWD increased survivorship of grass shrimp in laboratory and field experiments. These experimental results (1) support the hypothesis, commonly proposed but untested for freshwater habitats, that CWD can provide a refuge from predation for epibenthic fish and invertebrates and (2) extend the recognized functional importance of CWD in freshwater to estuarine and marine communities. We hypothesize that CWD is an especially important refuge habitat in the many estuarine and freshwater systems for which alternative physical structure (e.g., vegetation or oyster reefs) are absent or in low abundance.

Invasion pressure to a ballast-flooded estuary and an assessment of inoculant survival

The relationships between invasion pressure, post-transport inoculant survival, and regional susceptibility to invasion are poorly understood. In marine ecosystems, the movement and release of ballast water from ocean-going ships provides a model system by which to examine the interplay among these factors. One of the largest estuaries in North America, the Chesapeake Bay, receives tremendous amounts of foreign ballast water annually and thus should be at high invasion risk. To date, however, few introductions in Chesapeake Bay have been attributed to ballast release. To understand better the dynamics of this invasion process, we (1) characterized and quantified the biota arriving to Chesapeake Bay in foreign ballast water, (2) compared temperatures and salinities of ballast water and harbor water in upper Chesapeake Bay, and (3) tested experimentally survival of organisms collected from ballast water in temperatures and salinities characteristic of the region. From 1993 to 1994, we sampled planktonic and benthic organisms from 60 foreign vessels arriving to Chesapeake Bay. Our data show that the estuary is being inoculated by a diverse assemblage of aquatic organisms from around the world. Furthermore, the short transit time (≤15 d) for most vessels ensured that substantial numbers of larval and post-larval organisms were being deballasted alive. Most of the ballast water discharged into the upper Chesapeake Bay, however, was significantly higher in salinity (>20‰) than that of the receiving harbor. In laboratory tolerance experiments, ballast water organisms perished under such conditions. Thus, a mismatch in physical conditions between donor and receiver regions may explain the dearth of invasions in the upper Bay. It is likely that the lower Chesapeake Bay, which is more saline, remains at higher risk to ballast water invasion. Recognition of such intraregional differences should allow more focused predictions for monitoring and management.

Effect of an invasive crab upon a marine fishery: green crab, Carcinus maenas, predation upon a venerid clam, Katelysia scalarina, in Tasmania (Australia)

Abstract Nonindigenous species are increasingly recognized as altering marine and estuarine communities, causing significant changes in abundance and distribution of native species. Such effects are of particular concern to coastal fisheries. We experimentally determined the effect of the nonindigenous European green crab, Carcinus maenas, upon the stepped venerid clam, Katelysia scalarina, the basis for a fledgling clam fishery in Tasmania, Australia. First, we observed a trend of decreased juvenile (

The role of containerships as transfer mechanisms of marine biofouling species

Fouling of ships is an important historical and enduring transfer mechanism of marine nonindigenous species (NIS). Although containerships have risen to the forefront of global maritime shipping since the 1950s, few studies have directly sampled fouling communities on their submerged surfaces, and little is known about differences in the fouling characteristics among commercial ship types. Twenty-two in-service containerships at the Port of Oakland (San Francisco Bay, California) were sampled to test the hypothesis that the extent and taxonomic richness of fouling would be low on this type of ship, resulting from relatively fast speeds and short port durations. The data showed that the extent of macroorganisms (invertebrates and algae) was indeed low, especially across the large surface areas of the hull. Less than 1% of the exposed hull was colonized for all apart from one vessel. These ships had submerged surface areas of >7000 m2, and fouling coverage on this area was estimated to be <l7 m2 per vessel, with zero biota detected on the hulls of many vessels. The outlying smaller vessel (4465 m2) had an estimated coverage of 90% on the hull and also differed substantially from the other ships in terms of its recent voyage history, shorter voyage range and slower speeds. Despite the low extent of fouling, taxonomic richness was high among vessels. Consistent with recent studies, a wide range of organisms were concentrated at more protected and heterogeneous (non-hull) niche areas, including rudders, stern tubes and intake gratings. Green algae and barnacles were most frequently sampled among vessels, but hydroids, bryozoans, bivalves and ascidians were also recorded. One vessel had 20 different species in its fouling assemblage, including non-native species (already established in San Francisco Bay) and mobile species that were not detected in visual surveys. In contrast to other studies, dry dock block areas did not support many organisms, despite little antifouling deterrence in some cases. Comparisons with previous studies suggest that the accumulation of fouling on containerships may be lower than on other ship types (eg bulkers and general cargo vessels), but more data are needed to determine the hierarchy of factors contributing to differences in the extent of macrofouling and non-native species vector risks within the commercial fleet.