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Trends in Marine Biological Invasions at Local and Regional Scales: the Northeast Pacific Ocean As a Model System

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Trends in Marine Biological Invasions at Local and Regional Scales: the Northeast Pacific Ocean As a Model System Biological Invasions (2005) 7: 369–392 Ó Springer 2005 Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system Marjorie J. Wonham1,2,* & James T. Carlton3 1Department of Zoology, University of Washington, Box 351800, Seattle, WA 98195-1800, USA; 2Current address: Centre for Mathematical Biology, Department of Biological Sciences and Department of Mathematical & Statistical Sciences, University of Alberta, CAB 632, Edmonton, AB, Canada T6G 2G1; 3Maritime Studies Program, Williams College-Mystic Seaport, P.O. Box 6000, Mystic, CT 06355, USA; *Author for correspondence (e-mail: [email protected]; fax: +1-780-492-6826) Received 17 September 2003; accepted in revised form 8 March 2004 Key words: ballast water, Crassostrea, introduced species, invasion rate, invasion success, non-native species, Pacific Northwest Abstract Introduced species are an increasing agent of global change. Biogeographic comparisons of introduced biotas at regional and global scales can clarify trends in source regions, invasion pathways, sink regions, and survey effort. We identify the Northeast Pacific Ocean (NEP; northern California to British Colum- bia) as a model system for analyzing patterns of marine invasion success in cool temperate waters. We review literature and field surveys, documenting 123 introduced invertebrate, algal, fish, and vascular plant species in the NEP. Major invasion pathways were shipping (hull fouling, solid and water ballast; 1500s-present) and shellfish (particularly oysters) and finfish imports (commonest from the 1870s to mid- 1900s). The cumulative number of successful invasions over time increased at linear, quadratic, and exponential rates for different taxa, pathways, and regions within the NEP. Regional analysis of four major NEP estuaries showed that Puget Sound and the contiguous Straits had the most introduced spe- cies, followed by Humboldt Bay, Coos Bay and Willapa Bay. Data on cumulative shipping volumes pre- dicted smaller-scale, but not larger-scale spatial patterns in the number of shipping-mediated invasions. We identify the major challenges in scaling up from regional to global invasion analysis in cool temper- ate regions. Retrospective analyses for distinct biogeographic regions such as the NEP provide insight into vector dynamics and regional invasibility, and are a necessary foundation for monitoring and man- aging global change caused by biotic invasions. Introduction regional and global scales: this variation is the complex product of differences in the history, tim- Biological invasions are a major agent of global ing, and diversity of transport vectors, traits of the change (Ruiz et al. 1997; Vitousek et al. 1997; Sala introduced species, abiotic and biotic features of et al. 2000). In cool-temperate coastal communi- the invaded environment, and sampling strategies. ties, invasions by non-native marine and estuarine Biogeographic comparisons, based on a historical species are increasing around the world (Pollard and geographic synthesis of the introduced biota and Hutchings 1990a, b; Eno et al. 1997; Cohen within and between regions, can help elucidate the and Carlton 1998; Leppa¨koski and Olenin 2000; relative importance of these factors. Ruiz et al. 2000a; Levings et al. 2002). The number As a model system to analyze the growing and diversity of non-native species vary at both number of invasions in cool temperate marine 370 200 km Hecate Strait BRITISH Queen COLUMBIA Haida Charlotte Gwaii Sound (Queen Strait of Charlotte Georgia Islands) V an co uv er Vancouver Canada USA Strait of Juan de Fuca Puget Sound Seattle Pacific WASHINGTON Willapa Bay Ocean OREGON Coos Bay Humboldt Bay Cape Mendocino CALIFORNIA Figure 1. Major coastal water bodies of the cool temperate Northeast Pacific Ocean (Cape Mendocino, California, USA, to Haida Gwaii (Queen Charlotte Islands), British Columbia, Canada) treated in this review. ecosystems, we selected a 3000 km coastal ern limit of the Northern California Current Eco- biogeographic region in the Northeast Pacific region (OOPAC 1994), while the Queen Ocean (NEP) extending from Cape Mendocino, Charlottes represent the northern limit of contig- California (40°26.4¢ N, 124°24.3¢ W) to Haida uous coastal surveys before the considerably Gwaii (Queen Charlotte Islands), British Colum- more northern Alaskan datasets. We synthesized bia, (52–54° N, 131–133° W) (Figure 1 and records of introduced marine and estuarine spe- Table 1). Cape Mendocino represents the south- cies in this region and analyzed patterns in their Table 1. Regions within the Northeast Pacific treated in this analysis. Region Definition Abbreviation Puget Sound and Contiguous coastal waters of the Juan de Fuca Strait (entrance 48°23¢ N, SS Straits, WA and BC 124°44¢ W) and Puget Sound north to Hecate Strait (entrance 54°23¢ N, 132°26¢ W); includes Skeena, Fraser, Skagit, and Snohomish River estuaries and Duwamish waterway Coos Bay, OR Coos Bay (entrance 43°22¢ N, 124°22¢ W) including Coos River estuary CB Willapa Bay, WA Willapa Bay (entrance 46°37¢ N, 124°01¢ W) including Willapa, Palix and WB Naselle River estuaries Humboldt Bay, CA Humboldt Bay (entrance 40°23¢ N, 124°44¢ W) including Trinity, Eel, HB and Mad River estuaries Other Additional locales between Cape Mendocino and Haida Gwaii, including Other Yaquina Bay, OR, Columbia River Estuary, WA/OR, and Grays Harbor, WA) The total coastline treated here extends from Haida Gwaii, British Columbia (BC), Canada (54° N), through Washington (WA) and Oregon (OR) to Cape Mendocino, California (CA), USA (40° N). 371 taxonomic distribution, origins, invasion path- the late-1800s, iron-hulled ships coated with anti- ways, temporal trends, and regional distributions. fouling paint provided much less substrate for Lessons from regional-scale analysis of portions hull-fouling organisms. In lieu of solid ballast of the NEP highlight the challenges of making these ships carried water, and since its required global-scale comparisons. separation from oily bilge in the 1980s ballast water has served as a less polluted pathway for liv- Northeast Pacific invasion pathways ing organisms (Carlton 1985; Carlton and Geller 1993; Carlton and Hodder 1995). Although ship- Humans have inhabited the NEP coastline for ping tonnage is not necessarily a direct indicator thousands of years, but it was the 16th-century of number of ships or volume of ballast water, we arrival of European explorers and colonizers that use it as an indicator of the relative scale of ship- ushered in the present era of human-mediated ping pathways through time in the NEP. marine biological invasions. Dominant marine Shellfish imports to the NEP began in the late invasion pathways in the NEP have included ship- 1800s. The Atlantic oyster Crassostrea virginica ping (hull fouling, solid ballast, and water ballast) was imported from the 1880s to the 1930s. Paci- and imports for aquaculture and fisheries fic oysters, primarily C. gigas, were imported (reviewed by Carlton 1985, 1987; Hitchman 1990). from Japan beginning in the early 1900s and Shipping introductions presumably began with peaking around the 1950s; today experimental the arrival of European ships via the South imports are conducted only occasionally (D. Che- Atlantic and South Pacific in the mid-1500s, and ney, Director, Pacific Shellfish Institute, Olympia, Russian ships from the west and north Pacific in WA, USA, pers. comm.). Both oyster species the 1700s. Traffic increased during the gold rush served as invasion pathways for additional hitch- of the mid-1800s and again following the 1914 hiking invertebrates and algae (Quayle 1964, opening of the Panama Canal. In the US, 1988; Carlton 1979). Successful intentional intro- increased traffic was evident in the major ports of ductions have included several additional bivalve Puget Sound (Seattle and Tacoma); in ports of species; other molluscs and crustaceans were the outer coast, shipping remained at lower levels released but failed to establish (Carlton 1979). and in some areas has decreased in recent dec- Finfish were introduced intentionally to brackish ades (Figure 2). Early ships carried organisms on and coastal waters of the NEP in the late 1800s and in their wooden hulls and in solid ballast. By and early 1900s (Smith 1896; Fuller et al. 1999). 108 107 106 105 Shipping traffic (Short Tons) 104 1890 1910 1930 1950 1970 1990 2010 Year Figure 2. Annual shipping tonnage reported for selected years in selected Northeast Pacific ports: Humboldt Bay, California (solid squares), Coos Bay, Oregon (open triangles), Willapa Bay, Washington (open circles), and the Puget Sound ports of Seattle and Tacoma, Washington (solid diamonds). Note log scale. 372 Other potential invasion pathways that are less Atlantic salmon Salmo salar and the Mediterra- well documented include intentional and inadver- nean mussel Mytilus galloprovincialis). Although tent releases associated with the aquarium trade, the excluded species may in some cases play bait shipment, and planting and transplantation important ecological roles in marine and estuarine of eelgrass and salt marsh grasses. communities, our focus was on the changing com- position of established species living and repro- Materials and methods ducing in the these environments. In this review we synthesized current and histori- Data classification cal literature including peer-reviewed scientific publications, government reports, and unpub- Species were classified as native or introduced lished data. Field records from surveys
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